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Full text of "Bulletin of the Natural History Museum Entomology"

ISSN 0968-0454 



Bulletin of 
The Natural History Museum 



Entomology Series 




VOLUME 64 NUMBER 2 30 NOVEMBER 1995 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British 
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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Ent.) 

©The Natural History Museum, 1995 

Entomology Series 
ISSN 0968-0454 Vol. 64, No. 2, pp. 117-164 

The Natural History Museum 

Cromwell Road 

London SW7 5BD Issued 30 November 1995 

Typeset by Ann Buchan (Typesetters), Middlesex 
Printed in Great Britain at The Alden Press. Oxford 




Fig. 1 A. frontalis, habitus, cf • 



Bull. not. Hist. Mus. Lond. (Entomol.) 64(2): 117-163 Issued 30 November 1995 

Encyrtidae of Costa Rica 
(Hymenoptera: Chalcidoidea): the 
genus Aenasius Walker, 
parasitoids of mealybugs 
(Homoptera: Pseudococcidae) 

JOHN S. NOYES 

Department of Entomology, The Natural History Museum, Cromwell Road, 
London SW7 5BD 

H.REN 

Guangdong Entomological Institute, Guangdong, Guangzhou, P.R. China 

CONTENTS 



Synopsis 117 

I n t reduction 117 

Depositories 118 

Acknowledgements 119 

Aenasius Walker 119 

Systematic note 119 

Diagnostic characters 119 

Biology 119 

Use in biocontrol 120 

Distribution 120 

Identification of species 1 20 

Abbreviations used in text 121 

Key to Costa Rican species of Aenasius 121 

Review of species 123 

References 144 

Illustrations 146 

Index to scientific names 163 



Synopsis. The species of Aenasius from Costa Rica are revised. Of these, 16 are named 
and a further 20 recognised from males, but not named. Six species are described as new 
and 9 new specific synonymies are proposed. A dichotomous key to all species is 
provided and each species is further characterised by a diagnosis and notes are provided 
on their distribution and hosts. 



INTRODUCTION onomists in comparison with those of tem- 

perate areas such as Europe and North America. 

Early European taxonomists concentrated their 
Until recently, tropical parasitic Hymenoptera attention on their home ground because that is 
have received relatively little attention from tax- where it was easiest to obtain material on which 

©The Natural History Museum, 1995 



118 



J.S. NOYES ANDH. REN 



to work. The early work of North American 
taxonomists largely related to agriculture and the 
control of insect pests. Thus, it was not until 
relatively recently, with the use of modern col- 
lecting and preservation techniques, that the 
focus of these studies changed significantly. In 
particular, the use of Malaise traps greatly 
enhanced collection of parasitic Hymenoptera on 
a scale never before achieved. For instance, the 
work of Henry Townes revolutionised the study 
of Ichneumonidae by making available huge 
numbers of specimens collected from all over the 
world using his lightweight version of the Malaise 
trap (Townes, 1972). He also perfected a rapid 
technique for mounting specimens on the sides of 
pins. Although the use of a Malaise trap was seen 
as a valuable method for collecting microhy- 
menoptera, there was still one serious problem 
with using material collected by this method for 
taxonomic purposes. Small, weakly sclerotised 
specimens, such as Chalcidoidea, shrivelled 
badly when air-dried from alcohol. This made 
taxonomic work very difficult. The problem was 
overcome when it was realised that the use of 
critical-point drying could prevent specimens 
from collapsing (Gordh & Hall, 1976). This was 
an extremely important development in the study 
of weakly sclerotised microhymenoptera because 
it allowed specialists to remove specimens from 
alcohol and dry them without the danger of their 
integument collapsing. This revolutionised the 
study of smaller Chalcidoidea in tropical areas 
because it has allowed specialists to study a large 
amount of good-quality material from a wide 
area. 

The use of Malaise traps for collecting parasitic 
Hymenoptera has led to some surprising find- 
ings. Owen & Owen (1974) observed that Ich- 
neumonidae were no more diverse in tropical 
Africa than in temperate areas such as USA 
(Michigan) and Sweden. This was contrary to the 
generally accepted view that species richness, or 
diversity, increases with a decrease in latitude. 
Many plausible theories were put forward as to 
why at least some parasitic Hymenoptera in 
general follow this reverse trend. Yet findings for 
northern Sulawesi (Noyes, 1989) indicated 
strongly that whatever might be true for Ichneu- 
monidae did not seem to be the case for other 
groups of parasitic Hymenoptera, particularly 
the Chalcidoidea. The view that there is not an 
increase in species richness in Ichneumonidae in 
lower latitudes now seems to be well supported 
by some extensive studies in Costa Rica and 
eisewhere (Gauld, 1986; Gauld et al, 1992). 
These studies suggest that species richness in this 
group may be influenced by a number of factors 



including the possibility that many tropical 
insects may be unsuitable to act as hosts for 
ichneumonids because of an increase in toxins 
accrued by them from the plants upon which they 
feed ('poison host hypothesis'; Gauld et al., 
1992). This work has been supported by the 
extensive use of hundreds of Malaise traps to 
survey Ichneumonidae in Costa Rica since 1985. 
As a side-product of this survey, a vast quantity 
of microhymenoptera has become available for 
study. 

This unique collection has now made it pos- 
sible to survey in detail the microhymenopteran 
fauna of a tropical country for the first time. The 
results of the survey can be used for comparisons 
of species richness with similarly well-known 
temperate areas such as Great Britain. The col- 
lection can also be used for comparative studies 
on distribution within Costa Rica. Perhaps most 
important, by providing a sound framework for 
future work on this poorly studied group, taxo- 
nomic studies can ultimately provide the basis for 
future work on this group over a much wider 
area, perhaps the whole of South America. 

The present paper results directly from the 
above mentioned Malaise trap survey and from 
the collecting efforts of many other entomolo- 
gists too numerous to mention here. It is 
intended as the first contribution in a series of 
studies that will cover the taxonomy, distribution 
and known hosts of the 700 or so species of one 
family of Chalcidoidea, the Encyrtidae, that are 
known to occur in Costa Rica. It is hoped that 
these studies will encourage more detailed stud- 
ies of this family in Costa Rica and elsewhere in 
order to learn more about their biology, in 
particular their host ranges. Such work may 
facilitate the use of particular species in biologi- 
cal control programmes in other parts of the 
world should that become necessary. 

This first study is of the genus Aenasius, the 
species of which are relatively easy to recognise 
(see Fig. 1). The genus includes several species of 
actual or potential economic importance since 
their hosts are mealybugs (Homoptera: Pseudo- 
coccidae), many of which are important pests of 
agriculture in most parts of the world. 



DEPOSITORIES 



AMNH American Museum of Natural History. New 

York, USA 
BMNH The Natural History Museum, London, UK 
CNC Canadian National Collection. Ottawa, 

Canada 



ENCYRTIDAE OF COSTA RICA 



119 



IEE Instituto di Entomologfa Espariol, Madrid, 

Spain 

INBio Instituto National de Biodiversidad, Costa 
Rica 

MBA Museo Argentino de Ciencias Naturales. 
Buenos Aires, Argentina 

PPRI Plant Protection Research Institute, Preto- 
ria, South Africa 

TAMU Texas A&M University, College Station, 
Texas, USA 

USNM United States National Museum, Washing- 
ton D.C., USA 

ZISP Zoological Institute, Academy of Sciences, 
St Petersburg, Russia 



Acknowledgements. We thank The Royal Society, 
via the K.C. Wong fellowship, for allowing the junior 
author the opportunity to study at The Natural History 
Museum in London; his host institution, Guangdong 
Entomological Institute, permitted him one year's sab- 
batical leave. We also thank Doug Williams for his help 
in checking mealybug names. We especially thank Sue 
Lewis for preparing the electron micrographs (Figs 
86-97). Also thanks to Dr M.E. Schauff (USNM), Dr 
L. Masner (CNC) Dr J. Huber (CNC), Sa Izabel 
Izquierdo (IEE), Dr A. Bachman (MBA) and Dr E.L. 
Quinter (AMNH) for the loan or gift of material. Our 
special thanks to Pam Mitchell and Paul Hanson, for 
without their painstaking sorting of Malaise trap 
catches this project would not have been possible. 
Finally we would like to thank Aldo de Oyarzabal for 
the colour illustration of Aenasim frontalis (Fig. 1). 

Aenasius Walker 

Aenasius Walker, 1846: 180. Type species Aena- 
sius hyettus Walker, by monotypy. 

Neodiscodes Compere, 1931: 212-21 A. Type spe- 
cies Neodiscodes martinii Compere, by original 
designation and monotypy. Synonymised with 
Aenasius by Prinsloo, 1988: 1468. 

Pseudanasius Hayat, Alam & Agarwal, 1975: 
21-23. Type species Pseudanasius clavus 
Hayat, Alam & Agarwal, by original designa- 
tion and monotypy. Synonymised with Aena- 
sius by Hayat, 1981: 17. 

Systematic note 

Within the Encyrtidae, Aenasius belongs to the 
subfamily Tetracneminae, tribe Aenasiini. The 
tribal classification of the Tetracneminae has 
been reviewed by Noyes & Hayat (1994) with six 
tribes being recognised and diagnoses provided 
for the subfamily and each of the six included 
tribes. The genus has been diagnosed in a key to 
Neotropical encyrtid genera (Noyes, 1980), but 



can also be distinguised from all other genera of 
Encyrtidae using the characters given in the 
following diagnosis. 

Diagnostic characters 

Body generally squat and robust; mandibles 
bidentate with lower tooth short (Fig. 101), or 
with a short, upper, third tooth (Fig. 100). 
FEMALE: frontovertex (Figs 86-91) with con- 
spicuous piliferous punctures giving it the 
appearance of the surface of a thimble or golf 
ball; scrobes deep and frequently sharply mar- 
gined dorsally and laterally; scape varying from 
almost cylindrical (Fig. 18) to strongly broadened 
and flattened and less than twice as long as broad 
(Figs 3, 6); funicle 7-segmented (Figs 2-19), the 
first segment very small and normally visible only 
at higher magnifications on slide-mounted mate- 
rial; clava with sensory area enlarged, forming an 
oblique apical truncation which is longitudinally 
divided by a straight to sinuate line (Figs 9, 11, 
92, 93); forewings infuscate (Figs 20-45); apex of 
postmarginal and stigmal veins frequently con- 
nected by a naked, hyaline streak (Figs 32-45); 
gaster with hypopygium reaching apex; paraterg- 
ites sclerotised (Figs 46-49); last tergite more or 
less U-shaped; third valvulae usually free (Figs 
50, 52, 55). MALE: frontovertex either with 
piliferous punctures (Figs 96, 97) or with more 
irregular, sometimes coarse, sculpture (Figs 94, 
95); antennae either with 6 distinct funicle seg- 
ments and a small, entire clava (Figs 69-75), or 
with 2-5 anelliform segments and a relatively 
long, clava (Figs 76-83); wings hyaline (Figs 
62-68), rarely distinctly infuscate (Fig 65); post- 
marginal vein conspicuously longer than stigmal; 
phallobase (Figs 84, 85, 99) with a pair of distinct 
digiti, each with two or three apical hooks, 
outside each digitus a pair of bristles; aedeagus 
broad (Fig. 99) and about as long as mid tibial 
spur. 

Biology 

Where their hosts are known, all species are 
solitary endoparasitoids of mealybugs 
(Homoptera: Pseudococcidae). Little is known 
of their development but the egg and newly 
hatched larva have been described for Aenasius 
maplei Compere from North America (Maple, 
1947). In this species the deposited egg is typi- 
cally encyrtiform with the main part of the egg 
being attached to a smaller bulb by a narrow 
stalk. The deposited egg remains attached to the 
integument of the mealybug host via the stalk 
which, together with the collapsed bulb, projects 



120 



J.S. NOYES AND H. REN 



to the exterior of the host. The young larva has a 
single pair of functioning spiracles and remains 
attached to the remains of the egg shell and 
utilises atmospheric air via the protruding egg 
stalk and bulb. 

Use in biocontrol 

Although species of Aenasius must play an 
important role in the natural regulation of popu- 
lations of their mealybug hosts, there have been 
few attempts to utilise the species in biocontrol 
programmes (see Table 1). 

Distribution 

Of the 38 described species of Aenasius, 29 are 
known from the New World, eight are Afrotropi- 
cal and Oriental. The single remaining species, 
advena, is circumtropical, but of Neotropical 
origin. The Old World species form a group 
which is distinguished by the frontovertex being 
not more than one-sixth the head width, whilst in 
the New World species the frontovertex is con- 
spicuously wider, normally at least one-quarter 
head width. We have examined many unde- 
scribed species from South America. 

Within Costa Rica, species of Aenasius gener- 
ally tend to be most common in drier habitats at 
or below 500 m altitude, eg. Guanacaste, San 
Jose Province, and west of the central range. 
Malaise trap catches suggest that, although they 
are generally more common during the drier 
parts of the year (January to March in Guana- 
caste), they can also be common if there are 
longish dry spells during the wetter parts (eg. 
September-October in Guanacaste). This either 
reflects a real build-up in numbers due to the 
rapid population growth of their mealybug hosts, 
or greater parasitoid activity during drier peri- 
ods, or perhaps a combination of both. 



Identification of species 

Several keys have been published to the species: 
Compere (1937) for New World species, Kerrich 
(1967) for all known species (Old World species 
as Neodiscodes), Kaul & Agarwal (1985) for Old 
World species (as Neodiscodes) and Prinsloo 
(1988) for Afrotropical species. 

Some of these keys are based on characters 
which may be unreliable, e.g. coloration of 
antennal segments, or very small differences in 
the relative width of the scape or frontovertex. 
During our study we found that some characters 
appear to be reliable in defining species or 
groups of species. In the female these are: a) the 
presence or absence of a hyaline streak at the 
apex of the venation, b) the relative width and 
shape of the scape, c) the size and depth of the 
antennal scrobes, d) the appearance and distribu- 
tion of the piliferous punctures, e) the shape of 
the lower distal margin of the forewing, and f) 
the shape of the posterior margin of the last 
tergite of the gaster. In the male, four characters 
appear to be useful: a) the number and relative 
size of the funicle segments, b) the relative size of 
the clava, c) the sculpture of the frontovertex, 
and d) the presence or absence of particular 
sensilla on F6 or the clava (see Figs 102-105). 
The male genitalia of Aenasius do not seem to 
show a great deal of variation, although we have 
noted some variation in the relative size of the 
bristles at the bases of the digiti. However, we 
have not examined enough slide mounted mate- 
rial of any single species to determine the amount 
of intraspecific variation in this character. 

Although the present work deals with 16 
described species, we have recognised a further 
20 species from males collected in Costa Rica. As 
with most Encyrtidae, the taxonomy of Aenasius 
species is based entirely on female characters, 
and therefore these additional species are not 
being named in this paper, although some prob- 



Table 1 A summary of the use of Aenasius spp. in biological programmes worldwide (Abbreviations: E - 

established; NR - not released; ?? - no subsequent information; P - partial control; R - released, but no further 
information available; SC- successful control). 



Target pest species 


Aenasius sp. 


Country 


Year first used 


Result 


Source 


Phenacoccus herreni 


vexans 


Colombia 


1994 


SC 


J. Castillo (pers comm.) 


Dysmicoccus brevipes 


cariocus 


Hawaii 


1935 


99 


Swezey el al. (1939) 


Ferrisia virgata 


advena 


Hawaii 


1923. 1929. 1958 


P 


Bartlett in Clausen (1978). 
Funasaki, et al. (1988) 




advena 


California 


1966-1967 


R 


DeBach& Warner (1969) 


Phenacoccus manihoti 


phenacocci 


Africa 


1978 


NR? 


CIBC(1979, 1980) 


Pseudococcus tnaritimus 


paulistus 


USA 


9 


E? 


Compere (1937) 



ENCYRTIDAE OF COSTA RICA 

ably represent undescribed species. In order to 
facilitate their future recognition we include 
below short diagnoses of the males of all unasso- 
ciated species and include them in the key to 
species. 

Abbreviations used in text 



CL 
CW 
Fl,F2,etc 

EL 

EW 

FV 

FWL 

FWW 

GS 

HW 

HWL 

HWW 

MS 

MT 
OL 
OOL 



POL 

SL 
SW 



Length of costal cell of forewing 
Maximum width of costal cell of forewing 
First funicle segment, second funicle seg- 
ment, etc 

Maximum eye length 
Maximum eye width 
Minimum frontovertex width 
Forewing length 
Forewing width 

Maximum length of gonostylus (or third val- 
vula) 

Head width 
Hindwing length 
Hindwing width 

Malar space (the shortest distance from the 
eye to mouth margin) 
Mid tibia length 
Ovipositor length 

Ocular-ocellar line, or the shortest distance 
between each posterior ocellus and the adja- 
cent eye margin 

Posterior ocellar line, or the shortest dis- 
tance between the two posterior ocelli 
Scape length (excluding radicle) 
Maximum scape width 



Key to Costa Rican species of A enasius 

(Females and males) 

1 Clava three-segmented with an oblique apical trun- 
cation (Figs 2-5, 8-19, 98) never long and sausage- 
shaped). Females 2 

[Funicle 7-segmented, the first segment sometimes 
very small, anelliform and easily overlooked] 

- Clava entire, with apex rounded and frequently 
very long and sausage-shaped (Figs 69-83, 
102-105). Males 17 

[Funicle 3- to 6-segmented] 

2 Forewing without a naked, hyaline streak at apex of 
venation (Figs 20-31) 3 

- Forewing with naked, hyaline streak present (Figs 
1,32-45) 12 

3 Scape less than 2.5 x as long as broad (Figs 2, 8, 12, 
98) 4 

- Scape more than 2.5 x as long as broad (Figs 
3-15) 8 

4 Apex of last tergite broadly concave not with a 



121 

median incision (Figs 46, 47), either tegulae yellow 
or head and thorax with conspicuous silvery 
setae 5 

- Apex of last tergite with a median incision (Fig. 48, 
also as in Fig. 49), tegulae dark brown, head and 
thorax without conspicuous, lamellate silvery 
setae 6 

5 Head and thorax clothed in silvery setae, tegulae 
dark brown dives 

- Head and thorax clothed in brown setae, tegulae 
yellow vexans 

6 Funicle completely dark brown without yellow seg- 
ments phenacocci 

- Funicle with several yellow segments 7 

7 Scape mostly dark brown, piliferous punctures 
immediately above facial impression completely 
smooth and very shiny advena 

- Scape completely yellow, piliferous punctures 
immediately above facial impression dull lua 

8 Antenna almost completely yellow, only the pedicel 
dark brown cirrha 

- Antenna with scape and clava variously marked 
with brown 9 

9 Large piliferous punctures of frontovertex not, or 
hardly, extending between eyes and facial cavity; 
costal cell normally gradually tapering distally (Fig. 
27) rarely abruptly incised insularis 

- Large piliferous punctures extending at least half 
way between top of facial cavity and malar sulcus 
(as in Figs ,N(>-90); costal cell subrectangular, its 
apex abruptly incised at the point where the sub- 
marginal vein and marginal veins meet (Figs 29. 
31) 10 

10 Frontovertex more than one-third head width 
kerrichi 

- Frontovertex less than one-third head width ... 11 

11 Marginal vein about as long as stigmal vein (Fig. 
29) pelops 

- Marginal vein not more than two-thirds length of 
stigmal vein (Fig. 31) paulistus 

12 Scape not more than 2 x as long as broad, or hardly 
so (Figs 3, 4. 10) 13 

- Scape at least about 3 x as long as broad (Figs 
16-19) 15 

13 Scape about twice as long as broad and widest in 
middle with lower margin more or less evenly 
curved from apex to base (Fig. 18); funicle com- 
pletely black or with some yellow segments 
frontalis 

- Scape less than 1.9 x as long as broad and widest 
beyond middle, with lower margin distally bulging 



122 



J.S. NOYES ANDH. REN 



beyond insertion of pedicel and thus not evenly 
curved to base (Figs 3, 4, 6); funicle never with 
yellow segments, always completely black 14 

14 Facial impression acutely margined dorsally and 
relatively narrow and deep at this point, delimited 
by a concave or straight line (Fig. 88); dorsal 
surface of costal cell with setae relatively sparse, 
arranged in two or three lines (Fig. 34) bolowi 

- Facial impression not acutely margined dorsally and 
relatively broad and shallow at this point, delimited 
by a biconcave line (Fig. 89); dorsal surface of 
costal cell with setae dense and arranged in three or 
four lines (Fig. 38) caeruleus 

15 Lower part of apical margin of forewing straight or 
even slightly convex (Fig. 40); sensory area of clava 
not more than 2/3 as long as clava (both measured 
along longest axis) (Fig. 16); apex of last tergite 
without a median incision (as in Fig. 47) 
brasiliensis 

- Lower part of apical margin of forewing slightly 
emarginate (sometimes almost imperceptibly so) 
(Figs 42, 44); sensory area of clava more than 2/3 as 
long as clava (Fig. 19); apex of last tergite with a 
distinct median incision (as in Fig. 49) 16 

16 Scape longer than maximum width of eye; dorsum 
of thorax with a moderately strong blue-green lus- 
tre mitchellae 

- Scape slightly shorter than maximum eye width; 
dorsum of thorax dull, blackish, with a weak green- 
ish or purple lustre longiscapus 

17 Funicle composed of six distinct segments, none 
obscured by base of clava (Figs 69-75. 
102-105) 18 

- Funicle with fewer than six segments, these fre- 
quently hidden by base of clava, the latter some- 
times very long and sausage-shaped (Figs 
76-83) 30 

18 Funicle with both Fl and F2 relatively small, nei- 
ther with longitudinal sensilla and not wider than 
pedicel or hardly so; Fl not longer than pedicel 
(Figs 70, 72) 19 

- Funicle with at least F2 clearly wider than pedicel 
and usually with longitudinal sensilla; Fl frequently 
longer than pedicel (Figs 71, 73-75) 20 

19 Tegulae and alary sclerites dark brown, more or less 
concolorous with mesoscutum; forewings with post- 
marginal vein slightly longer than stigmal vein 
sp. A 

- Tegulae and alary sclerites orange brown, conspicu- 
ously paler than mesoscutum; forewings with post- 
marginal vein at least about 1.5 x as long as stigmal 
vein (Fig. 62) vexans 

20 Forewing with a naked, hyaline streak at the apex 
of the venation sp. B 



- Forewing without a naked, hyaline streak at the 
apex of the venation 21 

21 Pedicel not larger than Fl and usually shorter (Figs 
73-75, 104) 22 

- Either pedicel longer than Fl (measured dorsally or 
medially, excluding any ventral process, if present) 
or pedicel clearly larger than Fl (Figs 71, 102, 
105) 25 

22 Hind tarsi dark brown or dark orange-brown, as 
dark as hind tibiae or nearly so sp. C 

- Hind tarsi yellow to yellow-brown, clearly paler 
than hind tibiae 23 

23 Mesoscutum and scutellum with similar, fairly 
smooth and shallow imbricate-reticulate sculptu- 
re sp. D 

- At least scutellum with rough, reticulate, or 
punctate-reticulate sculpture 24 

24 Clava on external ventral surface near base, and 
usually F6, with an oval patch of specialised sensilla 
or setae which are clearly more dense than other 
setae present on these segments (Fig. 104) .. sp. E 

- Neither clava nor F6 with a similar patch of special- 
ised sensilla or setae sp. F 

25 Costal cell or forewing with at least three complete 
lines of setae dorsally (Fig. 64); clava with two oval 
sensory areas basally on outer ventral surface (Fig. 
102) bolowi 

- Costal cell of forewing with only two complete lines 
of setae dorsally, only occasionally with a few 
scattered setae comprising a partial third line; clava 
with at most only one oval sensory area basally . 26 

26 Antenna with only a very short, inconspicuous 
perpendicular seta on ventral surface of F6, this 
only about as long as adjacent recumbent setae and 
clearly less than one-third diameter of segment; 
forewings normally infuscate in basal half; larger 
species, generally >1 mm, smaller specimens 
rare brasiliensis 

- Antenna with a conspicuous perpendicular seta on 
ventral surface of F6 which is clearly much longer 
than the adjacent recumbent setae and at least 
nearly half as long as diameter of segment (Figs 103, 
105); forewings entirely hyaline; smaller species <1 
mm 27 

27 Frontovertex, below anterior ocellus, with regular, 
distinct piliferous punctures (as in Figs 96, 97) . 28 

- Frontovertex, below anterior ocellus, without pilif- 
erous punctures or if present than they are very 
indistinct and obscured by irregular, rough sculp- 
ture (as in Figs 94, 95) 29 

28 Antenna with ventral sensory areas on both F6 and 
clava; clava with a line of peg-like sensilla on inner 
surface extending from base to a little more than 



ENCYRTIDAE OF COSTA RICA 

one-third to apex (Fig. 103) sp. G 

- F6 and clava without any visible ventral sensory 
areas; clava with one or two peg-like sensilla about 
half way along ventral surface (Fig. 105) .... sp. H 

29 Clava basally with a single oval, ventral sensory 
area (similar to those in Fig. 102) and with a pair of 
basal peg-like sensilla on inner surface (similar to 
Fig. 105) sp. I 

- Clava without an oval, ventral sensory area near 
base and without peg-like sensilla on inner sur- 
face sp. J 

30 Funicle with more than two segments, distal seg- 
ments always obscured by base of clava, these 
sometimes only visible on slide-mounted specimens 
(Figs 77, 80, 81) 31 

- Funicle with two, clearly visible segments (if 
obscured dorsally by base of clava, then clearly 
visible ventrally) (Figs 78, 79, 82, 83) 35 

31 Frontovertex with distinct piliferous punctures 
which are smooth, or nearly so, and which give it 
the appearance of the surface of a thimble or golf 
ball (Fig. 97) 32 

- Frontovertex with piliferous punctures obscure, 
sculpture irregular and rough in appearance, not 
like the surface of a thimble or golf ball (Figs 94, 
95) 33 

32 Facial impression shallow and clearly remote from 
eye margins; piliferous punctures extending to 
nearly level with tops of antennal toruli sp. K 

- Facial impression relatively steep and more or less 
touching eye margins; piliferous punctures not 
extending between facial impression and eyes (Fig. 
97) phenacocciladvenal?longiscapus 

33 Clava at most about 1.5 x as long as scape .. sp. L 

- Clava at least about 2 x as long as scape (Figs 80, 
81) 34 

34 All tarsi dark brown and more or less concolorous 
with tibiae; funicle 5-segmented; clava about 4.5 x 
as long as broad (Fig. 80) sp. M 

- At least mid basitarsus yellow or testaceous-brown 
and conspicuously paler than mid tibia; funicle 
3-segmented; clava at least 7 x as long as broad 
(Fig. 81) sp. N 

35 Forewing hyaline 36 

- Forewing infuscate 38 

36 Clava about 3 x as long as scape and without a 
naked, ventral ridge (Fig. 79) sp. P 

- Clava not more than 2.5 x as long as scape and with 
a naked, longitudinal, ventral ridge (Fig. 76, 
78) 37 

37 Clava about 2 x as long as scape (Fig. 78); fron- 



123 

tovertex not more than one-third head width and at 
most about as wide as distance of anterior ocellus 
from facial impression sp. O 

- Clava about 2.5 x as long as scape (Fig. 76); 
frontovertex slightly more than one-third head 
width and nearly 1.5 x as broad as distance from 
anterior ocellus to facial impression paulistus 

38 Forewings strongly infuscate; clava with a conspicu- 
ous, naked ventral ridge extending along most of its 
length (Fig. 82) 39 

- Forewings only weakly infuscate; clava without a 
naked, ventral area (Fig. 83) 40 

39 Head about 2.5 x as wide as frontovertex; fron- 
tovertex about 1.5 x as wide as distance between 
anterior ocellus and facial impression sp. Q 

- Head about 2.25 x as wide as frontovertex; fron- 
tovertex a little more than twice as wide as distance 
between anterior ocellus and facial impression 
sp.R 

40 Antenna with Fl subquadrate, nearly as long as 
pedicel, clava about as long as scape sp. S 

- Antenna with Fl strongly transverse, clearly much 
shorter than pedicel; clava about 2 x as long as 
scape (Fig. 83) sp. T 



REVIEW OF SPECIES 



Aenasius dives sp. n. 

(Figs 2, 20, 21, 46, 52, 53, 56) 

Diagnosis. Female: head, dorsum of thorax 
and sides of gaster basally clothed in very con- 
spicuous whitish setae; antenna uniformly black; 
scape about 1.5 x as long as broad; sensory area 
of clava obliquely divided; frontovertex about 
one-quarter head width; piliferous punctures 
below anterior ocellus very shiny; scrobes deep, 
delimited dorsally and laterally; forewings with 
apex hyaline but with no hyaline streak present 
at apex of venation; lower part of outer margin of 
forewing slightly convex; scutellum with a deep, 
longitudinal median groove in basal half; apex of 
last tergite of gaster evenly and shallowly con- 
cave. 

Male. Unknown. 

Female. Length: 1.42-1.65 mm (holotype 1.62 
mm). Frontovertex metallic green, slightly cop- 
pery in ocellar area, below anterior ocellus pilif- 
erous punctures metallic green or blue-green, 
ridges between punctures purple, setae contrast- 
ing snow-white; temples and genae metallic 
green or blue green; facial impression metallic 



124 



J.S. NOYES ANDH. REN 



green mixed coppery, interantennal prominence 
mixed with purple; antennae black-brown; dor- 
sum of thorax dull dark blue, mixed with purple, 
almost black and clothed in very conspicuous 
pale brown, almost white setae; tegulae dark 
brown; propodeum medially blackish; mesopleu- 
ron blue-green; sides of propodeum coppery 
purple clothed in conspicuous pale setae; forew- 
ings (Fig. 20) more or less uniformly brown 
basally, but past apex of venation becoming 
gradually paler so that apical one-eighth or so is 
hyaline; no hyaline streak joining apices of post- 
marginal and stigmal veins (Fig. 21); hindwing 
hyaline; all coxae dark brown, fore coxae dis- 
tinctly metallic blue-green; femora and tibiae 
dark brown, but femora testaceous distally; fore 
tarsi almost completely brown, mid and hind 
tarsi yellow-testaceous, pretarsi dark brown; mid 
tibial spur dark brown; gaster dark purple-brown 
with conspicuous pale setae laterally at base. 

Head below anterior ocellus with large, con- 
spicuous, shiny piliferous punctures, each sepa- 
rated by a sharp ridge, punctures in ocellar area 
shallower less shiny, between antennal scrobes 
and anterior ocelli these punctures of slightly 
smaller diameter than anterior ocellus; two lines 
of punctures extending between facial impression 
and eyes nearly to level of lowest eye margin; 
scrobes delimited dorsally and at sides by a 
shallow carina; head in side view about twice as 
long as deep, evenly curved to top of scrobes, 
sides of facial impression slightly bulging out- 
wards and then abruptly angled inwards towards 
mouth; scape (Fig. 2) slightly bulging outwards at 
apex; clava slightly shorter than funicle, its apical 
sensory part about three times as long as ventral, 
straight margin of clava and divided by an 
oblique conspicuously sinuate line (as in Fig. 93); 
ocelli forming a distinctly acute. Relative mea- 
surements (holotype): HW 43.5; FV 10.5; EL 31; 
EW 21; MS 9; SL 24.5; SW 16; other proportions 
of antenna as in Fig. 2. 

Thorax without distinct piliferous punctures; 
sculpture on mesoscutum relatively deep, regu- 
lar, imbricate-reticulate, almost polygonally 
reticulate; scutellum with similar but slightly 
finer sculpture; scutellum about as long as broad 
and with a distinct marginal carina dorsally in 
apical one-fifth or so and with a deep, very 
conspicuous, longitudinal groove in its basal half; 
mid tibial spur clearly shorter than mid basitar- 
sus; fore wing with distribution of setae at base 
and proportions of venation as in Figs 20, 21; 
submarginal vein with a distinct subapical hyaline 
break; costal cell almost rectangular and con- 
spicuously incised at apex, dorsally with three or 
four lines of setae; no hyaline streak at apex of 



venation, lower part of apical margin slightly 
convex. Relative measurements (holotype): 
FWL 81, FWW 36, CL 36, CW 3; HWL 60, 
HWW 20. 

Gaster with last tergite apically slightly, but 
evenly concave (Fig. 46); ovipositor (Fig. 52, 53, 
56) with outer apical part of second valvifer 
strongly obliquely truncate and apically acute. 
Relative measurements (paratype): OL 37; GS 9; 
MT55. 

Male. Unknown. 

Variation. Very little of note in material avail- 
able. The frontovertex varies from slightly less 
than to more than one-quarter head width and 
the ovipositor varies from two-thirds to three- 
quarters as long as mid tibia. 

Hosts. Unknown. 

Distribution. Costa Rica. 

Material examined. 

Type material. Holotype $: COSTA RICA, 
Guanacaste Prov., Santa Rosa NP, Hacienda- 
3-0, 29.xi-20.xii. 1986 (Janzen & Gauld). 
Paratypes: COSTA RICA, 1$, Guanacaste 
Prov., Santa Rosa NP, Bosq. Hum.-ll-0, 
13.iv-4.v.l986 (Janzen & Gauld); 1$, Santa 
Rosa NP, Hacienda-3-0, 20.x. 1986-10.i. 1987 
(Janzen & Gauld); 2§, Guanacaste NP, Est. 
Mengo Vn Cacao, v. 1988 (Janzen & Gauld); 1$, 
Guanacaste NP, near HQ, 2-10.iii.1990 (J.S. 
Noyes); 1$, San Jose, Ciudad Colon. Hda El 
Rodeo, 16. ii. 1991 (Hym. Parataxonomists). 
Holotype and paratypes in BMNH, paratype in 
INBio. 

Comments. Aenasius dives is distinctive and can 
be separated from all other known species of the 
genus by the very conspicuous whitish setae on 
the head, dorsum of thorax and sides of propo- 
deum and gaster. 

Aenasius vexans Kerrich 

(Figs 22, 23, 47, 50, 51, 62, 70, 86, 98, 99, 101) 

Aenasius vexans Kerrich, 1967: 202-203. Holo- 
type 9> Brazil, USNM, examined. 
Aenasius tvexans Kerrich; Williams et al., 1981. 
Aenasius sp. nr vexans Kerrich; Lohr et al., 1990. 

Diagnosis. Female (length: 0.76-1.35 mm): 
head metallic green, dorsum of thorax similar, 
but duller; antenna more or less uniformly dark 
brown; tegulae orange basally; antenna (Fig. 98) 
with scape about 1.5 x as long as broad, sensory 
area of clava divided obliquely; head in side view 
about twice as long as deep and evenly curved to 



ENCYRTIDAE OF COSTA RICA 



125 



top of scrobes, sides of facial impression slightly 
bulging outwards and then abruptly angled 
inwards towards mouth; frontovertex (Fig. 86) 
about one-third to one-quarter head width, pilif- 
erous punctures below anterior ocellus shiny, 
scrobes moderately deep, not sharply delimited 
laterally; forewings (Fig. 22) apically hyaline 
with hyaline streak absent at apex of venation; 
postmarginal vein slightly longer than stigmal 
(Fig. 23); lower part of outer margin of forewing 
clearly convex; apex of last tergite of gaster 
almost straight, hardly concave (Fig. 47); ovi- 
positor as in Figs 50, 51. Male (length; 0.65-0.87 
mm): very similar to female but scape (Fig. 70) 
about 2.5 x as long as broad, all funicle segments 
transverse F4-6 subequal and each nearly twice 
as broad as long, clava about as long as F3-6 and 
without any conspicuous differentiated sensory 
areas or sensilla; forewings (Fig. 62) not or 
hardly infuscate; tarsi yellow; mid tibial spur 
yellow, occasionally pale brown; lateral bristle 
on phallobase (Fig. 99) about 0.25 x as long as 
aedeagus which is broad, spatulate and about 
one-third as long as mid tibia. 

Hosts. Recorded as a parasitoid of Phenacoccus 
herreni Cox & Williams (Williams et til. , 1981, as 
Aenasius Ivexans from yellow mealybug; Lohr et 
al., 1990), a pest of cassava in areas of Brazil and 
a potential pest of cassava in Colombia, Venezu- 
ela and the Guyanas (J. A. Castillo L., pers 
comm.). The eggs are laid into the 2nd and 3rd 
instar nymphs and adult females (Castillo L., 
pers comm.) 

Use in biocontrol. The species successfully 
controls Phenacoccus herreni Cox & Williams on 
cassava in Colombia (J. Castillo, pers. comm.). 

Distribution. Mexico, Costa Rica, Tobago, 
Trinidad, Colombia, Venezuela, Guyana, 
French Guiana, Ecuador, Peru, Brazil. 

Material examined. 

Type material. Holotype $: BRAZIL, Sao 
Paulo, ex Phenacoccus sp., xii.1935 (E. Hamble- 
ton) (USNM). Paratypes: MEXICO, 2$, 
Magdalena Is., Tres Marias, v. 1925 (H.H. 
Kiefer); BRAZIL, 1$, Sao Paulo, ex Phenacoc- 
cus No 3, xii.1935 (E. Hambleton) (BMNH). 

Non-type material: COSTA RICA, 6$, Gua- 
nacaste, Santa Rosa NP, various dates 
i.l986-iii.l987 (Janzen & Gauld); TOBAGO, 
19, St Paul, Pamatuvier Valley, edge of rainfor- 
est, 20.vii.1976 (J.S. Noyes); TRINIDAD, 1$, 
I.C.T.A., ex Phenacoccus hibsici on Hibiscus, 
No 25, v.1953 (F.D. Bennett); 29, Curepe, 
Santa Margarita Circular Road, 26.ix-26.x.l974 



(M.N. Beg); 5$, St George, various localities 
and dates, vi— vii. 1976 (J.S. Noyes); 5$, Nariva, 
Cocos Bay, coastland and mangrove swamp, 
28. vii. 1976 (J.S. Noyes); 1$, St. Augustine, ex 
mealybug on Lantana montividensis, V.1982 
(M.W.J. Cock); 6$, Curepe, ex Phenacoccus sp. 
on tomato, ix.1983 (F.D. Bennett); COLOM- 
BIA, 339, 26cf, Cali, CIAT, lab culture from 
Venezuela, Bolivia Sta (Upsta) and Sucre Sta 
(Ayacucho and Cumana), ex Phenacoccus her- 
reni on cassava, xi.1991 (J. Castillo); VENEZU- 
ELA, 119. UO\ Sucre, ex Phenacoccus herreni 
15. vi. 1989 (J. Castillo); GUYANA, 19, 1Q\ 
Enmore Est., x.1979, ex Phenacocci herreni (as 
P. manihoti misident.) on cassava, x.1979 (F.D. 
Bennett); 19- Enmore Est., ex Phenacoccus on 
cassava, 11. xi. 1977 (M. Yaseen); Id", Dalgin, ex 
Phenacocci herreni (as P. manihoti misident.) on 
cassava, xi. 1978 (M. Yaseen); ld\ Diamond, ex 
Phenacocci herreni (as P. manihoti misident.) on 
cassava, xi.1979 (M. Yaseen); FRENCH GUI- 
ANA, 19. Km 30 Highway, ex Phenacocci her- 
reni (as P. manihoti misident.) on cassava, 
xi. 1978 (M. Yaseen); 39- Sinnamary, ex Phen- 
acocci herreni (as P. manihoti misident.) on 
cassava, xi. 1978 (M. Yaseen); 29. 5d\ Mana, ex 
Phenacocci herreni (as P. manihoti misident.) on 
cassava, xi.1978 (M. Yaseen); ECUADOR, 19, 
Pichincha, Tinlandia, 800m, 2.ii.l983 (Masner & 
Sharkey); PERU, Cuaco, Quilambamba, 
24-26.xii.l983(L. Huggert); 1$, Madre de Dios, 
Pto Maldonado, 3. i. 1984 (L. Huggert); BRA- 
ZIL, 39, 2d", Santa Lucia, xi.1977, ex Phenacoc- 
cus on cassava, xi,1977 (M. Yaseen); 49. 30\ 
Macapa, ex Phenacoccus on cassava, 22. xi.1977 
(M. Yaseen); 29. Porto Grande, ex Phenacoccus 
on cassava, 24.xi.1979 (M. Yaseen); 69. 2o\ 
Para, Alanquer, ex Phenacoccus herreni on cas- 
sava, 11.x. 1985 (B. Lohr). Material in BMNH, 
CNC, USNM, TAMU, ZISP, PPRI, INBio. 

Comments. Aenasius vexans can be mistaken 
for phenacocci but is separated on the coloration 
of tegulae and relative length of postmarginal 
vein (see comments under phenacocci). 

Aenasius phenacocci Bennett 

(Figs 8, 9, 24, 25,48, 55,63) 

Aenasius phenacocci Bennett, 1957: 569-570. 
Holotype 9- Trinidad, USNM, examined. 

Aenasius flandersi Kerrich, 1967: 204, 221. Holo- 
type 9. USA, USNM, examined, syn. n. 

Diagnosis. Female (length: 0.88-1.84 mm): 
head mostly metallic green or blue green; 
antenna uniformly black brown; pronotum and 



126 



J.S. NOYES ANDH. REN 



mesoscutum weakly metallic green or blue- 
green; scutellum with a weak purple or blue- 
green lustre; tegulae dark brown; antenna (Fig. 
8) with scape about 1.6 x as long as broad; 
sensory area of clava divided obliquely into 
almost equal portions (Fig. 9); head in side view 
very slightly more than twice as long as deep; 
slightly curved above scrobes, below scrobes 
more strongly curved towards mouth; frontover- 
tex about one-third (small specimens) to one- 
fifth (large specimens) head width, piliferous 
punctures below anterior ocellus shiny, scrobes 
moderately deep, not sharply delimited; forew- 
ings (Fig. 24) apically hyaline without a hyaline 
streak at apex of venation; postmarginal vein not 
quite reaching level with apex of stigmal (Fig. 
25); lower part of outer margin of forewing very 
slightly convex; apex of last tergite of gaster 
inconspicuously medially incised (Fig. 48); ovi- 
positor (Fig. 55) about as long as mid tibia. Male: 
indistinguishable from that of advena Compere 
(see diagnosis); forewing hyaline (Fig. 63). 

Hosts. Recorded as a parasitoid of Phenacoccus 
madeirensis Green (as P. gossypii, probable misi- 
dentification, D. Williams, pers. comm.) (Ben- 
nett, 1957) and from the same host on Acalypha 
sp., Hibiscus sp. and an unidentified mealybug 
on Pittosporum and cotton (Kerrich, 1967). Also 
recorded from Phenacoccus herreni Cox & Will- 
iams (Lohr et al., 1990). The records below from 
P. gossypii Townsend & Cockerell on Lantana 
montividensis , Phenacoccus grenadensis Green & 
Laing on Cordia curassavica and Tussacia sp. are 
probably all misidentifications of P. madeirensis 
(D. Williams, pers. comm.). Also noted here as a 
parasitoid of Ferrisia virgata (Cockerell) on Sida 
sp. 

Distribution. USA (California), Costa Rica, 
Cayman Islands, St Vincent, Trinidad, Colom- 
bia, Guyana, French Guiana, Ecuador, Peru, 
Uruguay. 

Material examined. 

Type material. Holotype $ of Aenasius phen- 
acocci, TRINIDAD, I.C.T.A., ex Phenacoccus 
gossypii on Acalypha, v. 1955 (F.D. Bennett) 
(USNM Type No 63501). Paratypes of Aenasius 
phenacocci: TRINIDAD, 2$, I.C.T.A., St. 
Augustine, ex Phenacoccus gossypii on Aca- 
lypha, vi.1955 (F.D. Bennett) (in BMNH). Holo- 
type $ of Aenasius flandersi: USA, California, 
San Diego, Balboa Park, on Pittosporum, 
15.viii.1958 (S. Flanders) (Type No 2122 
USNM). 

Non-type material. COSTA RICA, 1$, Gua- 
nacaste, Santa Rosa NP, Hacienda 1-0, 



14.viii-6.ix.1986 (Janzen & Gauld); 1$, San 
Jose, Ciudad Colon, 800m, iii-iv.1990 (L. 
Fournier); CAYMAN ISLANDS, Grand Cay- 
man, West Bay, Willie Farringdon Drive, ex 
Ferrisia virgata on Sida, 16.x. 1987 (F.D. Ben- 
nett); ST VINCENT, 1$, 207, West Indies 
99-331, Aenasius hyettus Wlk. How.(H.H. 
Smith); TRINIDAD, 4$, I.C.T.A., ex Phen- 
acoccus gossypii on Hibiscus, v-vi.1953 (F.D. 
Bennett); 15$, St. Augustine, ex Phenacoccus 
gossypii on Acalypha, v. 1955 (F.D. Bennett); 
2$, lcf, St. Augustine, ex mealybugs, hi. 1961 
(F.D. Bennett); 2$, Curepe, Sta. Margarita, 
26.ix-26.x.l974 (M.N. Berg); 1$, St. George, 
San Juan, ex mealybug; 18.vii.1976 (F.D. Ben- 
nett); 1$, St Augustine, Malaise trap; ix.1976 
(F.D. Bennett); 11$, St George, various locali- 
ties, vi-viii. 1976 (J.S. Noyes, F.D. Bennett); 1$, 
St. Andrew, Oropuche, cocoa and banana plan- 
tations, 28.V.1976 (J.S. Noyes); 3$, CIBC lab 
culture ex Phenacoccus grenadensis, 1977 (M. 
Yaseen); 1$, Maracas Valley, xii.1977 (F.D. 
Bennett); 3$, Curepe, ex Phenacoccus gossypii 
on Lantana montividensis, iv.1977 (F.D. Ben- 
nett); 5$, Curepe, ex Phenacoccus grenadensis 
on Cordia curassavica, iii.1979 (F.D. Bennett); 
2$, Curepe, CIBC lab culture on Phenacoccus 
grenadensis, iv.1979 (F.D. BEnnett); ld\ ex 
Phencacoccus grenadensis on Tussacia, x.1979 
(F.D. Bennett); 1$, St George, Arima Valley 
(1st mile), edge of rainforest, 3.vii. 1976 (J.S. 
Noyes); 1 gynandromorph, St George, St Aus- 
gustine, malaise trap, 15.vii-13.viii.1976 (J.S. 
Noyes); GUYANA, 1$, Georgetown, on orna- 
mentals; 15.x. 1961 (F.D. Bennett); 4$, Enmore, 
ex Phenacoccus Imanihoti on cassava, x.1979 
(M. Yaseen); COLOMBIA, 2$, Cali, CIAT, ex 
Phenacoccus grenadensis on cassava, xi.1977 
(F.D. Bennett); 1$, Palmira, ex Phenacoccus on 
Acaplypha, 14. xi. 1979 (F.D. Bennett); ECUA- 
DOR, 1$, Rio Palenque, forest, 4.ii.l983 (L. 
Huggert). Material in BMNH, INBio. 

Comments. We have been unable to find any 
significant differences between the type material 
of flandersi examined and the series of phen- 
acocci listed above. We therefore treat the two 
names as synonymous. A. phenacocci is 
extremely close to advena differing only in the 
colour of the funicle segments, other differences 
given by Kerrich (1967) are seemingly unreliable. 
For the present, we continue to treat the two 
species as valid until further material becomes 
available. 

Superficially, Aenasius phenacocci may be mis- 
taken for vexans, but is distinguished on the 
completely brown tegulae and postmarginal vein 



ENCYRTIDAE OF COSTA RICA 



127 



of forewing not being longer than the stigmal. In 
vexans the tegulae are largely yellow and the 
postmarginal vein is slightly longer than the 
stigmal. 

Aenasius advena Compere 

(Figs 12, 77, 84, 97) 

Aenasius advena Compere, 1937: 384, 388-389, 
393. Holotype 9- Hawaii, USNM, examined. 

Aenasius ianthinus Compere, 1937: 388, 391, 
393-394. Holotype $, Panama, USNM, not 
examined, syn. n. 

Pseudanasius clavus Hayat, Alam & Agarwal, 
1975: 23-24. Holotype, gynandromorph, 
India, BMNH, examined. Synonymised with 
advena by Hayat, 1981. 

Diagnosis. Female (length: 1.14-2.16 mm): as 
for phenacocci but antenna with at least some 
funicle segments yellow (Fig. 12) (see also discus- 
sion below); ovipositor generally about one-third 
longer than mid tibia. Male (length: 0.83-1.40 
mm): duller than female; forewings hyaline; tarsi 
yellow, mid tibial spur dark brown; scape (Fig. 
77) about 2.5 x as long as broad and widest in 
middle, funicle with 5 anelliform segments, distal 
3 or 4 enclosed by base of clava which is about 
7-8 x as long as broad, 3-3.5 x as long as scape 
and without a longitudinal naked ventral ridge; 
frontovertex (Fig. 97) about one-third head 
width and with distinct piliferous punctures 
which do not extend between eyes and facial 
impression, facial impression touching eyes; dis- 
tance from anterior ocellus to facial impression 
about 0.6 x minimum width of frontovertex; 
genitalia as in Fig. 84. 

Hosts. Aenasius advena has been recorded 
throughout its range as a parasitoid of Ferrisia 
virgata (Cockerell) on a variety of plants (Ben- 
nett, 1957; Compere, 1937; Kerrich, 1967 
DeBach & Warner, 1969; Shafee et al., 1975 
Hayat et al., 1975; Prinsloo & Annecke, 1979 
De Santis, 1980), also from Brazil as a parasitoid 
of Pseudococcus longispinus (Targioni-Tozzetti) 
(De Santis, 1980) and Pseudococcus sp. on Mac- 
quilla tomentosa (Kerrich, 1967). 

Use in biocontrol. Aenasius advena has been 
introduced into Hawaii (Bartlett in Clausen, 
1978; Funasaki et al., 1988) and California in 
1966-1967 (DeBach & Warner, 1969) for the 
control of Ferrisia virgata, a mealybug pest of 
ornamentals and other plants. It has been par- 
tially successful in controlling this pest in Hawaii. 

Distribution. A circumtropical species found 



throughout most of the Neotropical, Afrotropical 
and Oriental regions (see below and above refer- 
ences). 

Material examined. 

Type material. Holotype 9 of Aenasius advena: 
HAWAII, Oahu, Flates west of Makapuu Head, 
on wild cotton, ll.ii.1934 (O.H. Swezey) (Type 
No 51984 USNM). Paratypes of Aenasius 
advena: HAWAII, 2$, 2d", Oahu, Koko Head, 
15.iii, 13. ii, 20.ii, 27. ii. 1934 (O.H. Swezey). 
Holotype (gynandromorph) of Pseudanasius cla- 
vus, INDIA, Tamil Nadu, Shencottah, ex Ferri- 
sia virgata, 6.iii.l967 (M. Hayat) (BMNH). 
Paratype of Pseudanasius clavus: INDIA, 10". 
Tamil Nadu, Shencottah, ex Ferrisia virgata, 
16.iii.1967 (M. Hayat) (BMNH). 

Non-type material. USA, 1$, Florida, Key 
Largo, 26.xii.1954 (H.V. Weems jr); COSTA 
RICA, 19, Guanacaste Prov., Finca Taboga, 
6mi S, 6mi W. Canas, 10°19'N 89°09'W, ii.1967 
(H.A. Hespenheide); 19. Guanacaste Prov., 
Santa Rosa NP, Hacienda 2-C, 22.vi-13.vii.1985 
(Janzen, Gauld); 4cf, Gunacaste Prov., Santa 
Rosa NP, Hacienda 3-0, 14.viii-18.x. 1986 (Jan- 
zen & Gauld); PANAMA, 1$, Aquadulce, 
27.vii.1976 (F.D. Bennett); TRINIDAD, 1$, 
Manzanilla, ex Ferrisia on coconut, v. 1952 (F.D. 
Bennett); 1$, No. 1 (Bennett); 39, 1.C.T.A., ex 
Ferrisia on Gliricidia, viii.1953 (F.D. Bennett); 
VENEZUELA, 19, Bahia del Mar. ex Ferrisia 
virgata on ornamental, 7.xii. 1981 (M. Yaseen); 
BRAZIL, 19. Rio de Janeiro, on citrus; 
27.x. 1962 (A. Perachi); 1$, Sao Paulo, Araras, 
on citrus, 27. iv. 1984 (F.D. Bennett); BOLIVIA, 
29, La Cororada, ex Phenacoccus manihoti, 
20.vi.1982 (M. Yaseen); P.R.CONGO, 19, 3d\ 
Brazzaville, ex Saissetia oleae on cassava, 1987 
(A. Biassangama); INDIA, 19, 20\ Delhi, 
IARI area, x.1979 (Z. Boucek); 29, Gujarat, 
Jungadh, ex mealybug in croton, 28.viii.1988 
(V.N. Patel); 1$, 2d\ Madhya Pradesh, Jabal- 
apur, ex Ferrisia virgata 25. xi. 1966 (B.N. Modi); 
59, Karnataka, Bangalore, ex Ferrisia virgata on 
Thervetia nervifolia, 2.iv.l982; 29, Karnataka, 
Bangalore, Hessaraghatta, ex Planococcus citri, 
iii.1989; 69, Karnataka, Shanthigadu, ex Ferrisia 
virgata on cashew, iv.1990 (Sundararaju); 39, 
3d", Karnataka, Theerthalli, Bakalapura Est., ex 
Ferrisia virgata on coffee, v. 1990 (P.K. Bhat); 
39, 60", Karnataka, Netragonda Est., ex Ferrisia 
virgata on coffee, v. 1990 (Sreedharan, Balakrish- 
nan); 19, Tamil Nadu, Coimbatore, 
25. ix-1. x.1979 (J.S. Noyes); 1$, Tamil Nadu, 
3Km E. Manjaler Dam, 15-18.X.1979 (J.S. 
Noyes); 39, lcf, Kerala, Wynad, ex Ferrisia 
virgata on robusta coffee, 15. v. 1987 (C. 



128 



J.S. NOYES ANDH. REN 



Prakasan); ld\ Kerala, ex Ferrisia virgata on 
cashew, 17. xi. 1989 (Fathima); 4$, Andaman 
Islands, Port Blair, ex mealybug on citrus, 
22.iii.1988 (B.S. Bhumannaver); BANG- 
LADESH, 1$, Dacca, ex Pseudococcus virgatus, 
1963 (Gov't Entom.); THAILAND, 2$, 2cf, 
Bangkok, on Samanea saman, 1983 (S. 
Boonkong); MALAYSIA, 2$, 2cf, Selangor, 
RRI, 3222-2-58, CIE Coll. No 16100; 2$, lcf, 
Selangor, Bukit Rotan, Sg. Buloh Estate, 
16.viii.1958; PHILIPPINES, 1$, Manila, guava, 
xii.1958 (Krauss); FIJI, 6$, 2c? , Naduruloulou, 
ii,1949, ex mealybug on twigs oiAlbizzia lebbeck 
(B.A. O.Connor); HAWAII, 1$, Johnston 
Island, 3508, on Virex trifolia, 8.vii.l948 (L.B. 
Loring). Material in BMNH 

Comments. Aenasius advena is close to phen- 
acocci and lua (see comments under those spe- 
cies). 

Aenasius ianthinus was treated as a variety of 
advena by Kerrich (1967) and therefore we 
regard it here as a synonym of advena. 

Aenasius lua sp. n. 

Diagnosis. Female: antenna with scape and 
funicle segments yellow; scape about twice as 
long as broad; sensory area of clava divided into 
two unequal parts by a sigmoidal, oblique line; 
frontovertex about one-fifth head width; pilifer- 
ous punctures below anterior ocellus dull; 
scrobes not deep and not sharply delimited; 
forewings infuscate but with apex hyaline and 
without a naked hyaline streak at apex of vena- 
tion; lower part of outer margin of forewing not 
emarginate; apex of last tergite of gaster with a 
slight median incision. Male: unknown. 

Female. Length: 1.73 mm. Head metallic 
green, ocellar area and above facial impression 
slightly coppery; mouth margin, lower genae and 
temples dark coppery-purple; radicle dark brown 
and pedicel dark brown, scape and funicle yel- 
low, clava (except sensory area) brown; setae on 
frontovertex pale translucent brown; dorsum of 
thorax blackish much less shiny than frontover- 
tex, with only a slight purplish sheen; setae 
translucent brown; tegulae dark brown; sides of 
thorax dark brown; forewings more or less uni- 
formly brown in basal half, apical one-eighth or 
so hyaline; hyaline streak at apex of venation 
absent; hindwing hyaline; legs, except tarsi, dark 
brown; mid tibial spur dark brown; tarsi whitish 
or pale yellow, pretarsi brown; gaster dark 
brown with an inconspicuous purple sheen. 

Head with conspicuous piliferous punctures, 
each about the same diameter as an ocellus, 



those on frontovertex relatively dull with micror- 
idges radiating out from the bases of the setae, 
between eyes and facial impression quite shiny; 
three lines of piliferous punctures present 
between facial impression and each eye, these 
nearly reaching malar sulcus; facial impression 
not deep and not delimited by sharp carinae, 
either dorsally or laterally; head in side view 
almost twice as long as deep, almost flat above 
scrobes, but below this distinctly curved towards 
mouth; scape with lamina not bulging outwards 
at apex; clava about as long as funicle and pedicel 
together, its apical sensory part about 0.8 x as 
long as clava and divided by an oblique line into 
nearly equal parts; ocelli forming an acute, 
almost equilateral, triangle. Relative measure- 
ments: HW 127; FV 26; EL 83; EW 62; MS 36; 
SL 45; SW 20; other proportions of antenna 
similar to advena (Fig. 12). 

Piliferous punctures on mesoscutum small and 
shallow, separated by more than their own diam- 
eters, those on scutellum more indistinct; sculp- 
ture on mesoscutum shallow, regular imbricate- 
reticulate, nearly polygonally reticulate 
posteriorly; scutellum with similar sculpture; 
scutellum about as long as broad and with a 
distinct marginal carina dorsally in apical one- 
fifth or so; mid tibial spur about as long as mid 
basitarsus; forewing with distribution of setae at 
base and proportions of venation similar to phen- 
acocci (Fig. 24), but costal cell with only two 
lines of setae dorsally; hyaline streak absent at 
apex of venation, lower part of apical margin of 
forewing straight or even slightly convex. Rela- 
tive measurements (holotype): FWL 195, FWW 
91;HWL141,HWW55. 

Gaster with last tergite hardly concave but 
with a slight median incision; ovipositor at least a 
little longer than the mid tibia. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Costa Rica. 

Material examined. 

Holotype $: COSTA RICA, Puntarenas, Mon- 

teverde, 15-16.vii.1988 (L. Masner) (CNC). 

Comments. Aenasius lua is very close to 
advena, females of both species having a rela- 
tively broad scape, dark tegulae, lack of a hyaline 
streak at apex of venation and the posterior 
margin of the last abdominal tergite with a 
median notch. The species can be separated on 
the sculpture of frontovertex, coloration of scape 
and density of setae on dorsal surface of costal 
cell. A. lua has the scape completely yellow, the 



ENCYRTIDAE OF COSTA RICA 



129 



piliferous punctures of the frontovertex dull and 
only two lines of setae on the dorsal surface of 
the costal cell whereas in advena the scape is 
mostly dark brown, the piliferous punctures of 
the frontovertex are very shiny and, in similar- 
sized specimens, the costal cell has at least three 
lines of setae dorsally. 

Aenasius cirrha sp. n. 

Diagnosis. Female: antenna yellow with 
pedicel dark brown; scape about three times as 
long as broad; sensory area of clava not conspicu- 
ously divided, division in apical half and longitu- 
dinal; frontovertex a little over one-fifth head 
width; piliferous punctures below anterior ocel- 
lus dull; scrobes moderately deep, not sharply 
delimited laterally; forewings infuscate proxi- 
mally, slightly less than apical one-third hyaline; 
apex of venation without a naked, hyaline streak; 
costal cell gradually tapering to its apex, but with 
a slight apical incision before the marginal vein; 
lower part of outer margin of forewing slightly 
convex; apex of last tergite of gaster evenly and 
shallowly concave, slightly incised medially. 

Female. Length: 1.62 mm. Ocellar area and 
area above facial impression metallic coppery 
and purple; temples, scrobes and genae metallic 
green with a coppery sheen; interantennal promi- 
nence and mouth margin quite strongly coppery 
purple; piliferous punctures between anterior 
ocellus and facial impression rather dull; setae on 
frontovertex translucent brown; antenna with 
radicle orange-brown, pedicel dark brown dor- 
sally, the remainder yellow; dorsum of thorax 
blackish with a slight purple and brassy sheen; 
setae dark brown; tegulae dark brown; sides of 
thorax brown; forewings brown proximally, but 
gradually fading distally so that a little less than 
apical one-third is hyaline; apex of venation 
without a naked, hyaline streak; hindwing hya- 
line; all coxae dark brown; femora dark brown, 
paler apically, especially hind femora; fore and 
mid tibia brown with apices yellowish; hind tibia 
mostly yellow, slightly infuscate along dorsal 
margin proximally; tarsi pale yellow, mid tarsi 
slightly darker; mid tibial spur dark brown; 
gaster dark brown with an inconspicuous blue or 
purple sheen. 

Head with conspicuous, dull piliferous punc- 
tures, each separated by a sharp ridge, punctures 
in ocellar area shallower less shiny, between 
facial impression and anterior ocellus these punc- 
tures slightly smaller than ocelli; punctures not 
extending between eyes and facial impression; 
scrobes sharply delimited dorsally but not later- 



ally; head in side view about twice as long as 
deep, slightly convex above facial impression; 
more strongly curved towards mouth about level 
with top of impression; clava at least slightly 
longer than funicle and pedicel together, its 
apical sensory part about two-thirds as long as 
clava and not conspicuously divided, the division 
only slightly oblique and almost entirely in apical 
half; ocelli forming a hardly acute angle. Relative 
measurements: HW 125; FV 29; EL 84; EW 61; 
MS 30; SL 46; SW 15; other proportions of 
antenna similar to insularis (Fig. 13). 

Thorax with small, indistinct piliferous punc- 
tures; sculpture on mesoscutum regular 
imbricate-reticulate; scutellum similar but sculp- 
ture more reticulate; scutellum about as long as 
broad and with a distinct marginal carina dorsally 
in apical one-fifth or so; mid tibial spur about as 
long as mid basitarsus; forewing with distribution 
of setae at base and proportions of venation 
similar to those of pelops (Figs 28, 29), postmar- 
ginal vein about as long as stigmal; submarginal 
vein with a distinct subapical hyaline break; 
costal cell slightly more than 6 x as long as 
broad, gradually tapering apically, but with apex 
slightly incised before marginal vein, dorsally 
with two or three lines of setae; apex of venation 
without naked, hyaline streak, lower part of 
apical margin slightly convex. Relative measure- 
ments (holotype): FWL 226. FWW 98, CL 85, 
CW 13.5; HWL 146, HWW 55. 

Gaster with last tergite apically biconvex either 
side of a median incision. 

Hosts. Unknown. 

Distribution. Costa Rica. 

Material examined. 

Holotype $, COSTA RICA, Puntarenas, Mon- 

teverde, 15-16. vii. 1986 (L. Masner). In CNC. 

Comments. Aenasius cirrha is close to paulistus 
and insularis, all three species lacking the hyaline 
streak at the apex of the venation, having a scape 
more than 2.5 x as long as broad and costal cell 
gradually tapering in apical half or so. A. cirrha 
can be separated from these species by the 
almost entirely yellow antennae, the apex of the 
costal cell slightly but distinctly incised, whilst in 
the other named species the scape and clava are 
largely brown and the costal cell more or less 
gradually tapers at this point. In addition, it can 
be separated from paulistus by the piliferous 
punctures not extending between the eyes and 
facial impression. 



130 



J.S. NOYES ANDH. REN 



Aenasius insularis Compere 

(Figs 13, 26, 27, 49, 54, 58) 

Aenasius insularis Compere, 1937: 392, 400-401. 
Holotype $, Mexico, USNM, examined. 

Diagnosis. Female (length 1.11-2.03 mm): as 
for cirrha but frontovertex generally metallic 
green; antenna (Fig. 13) with scape marked 
variously with brown proximally; pedicel, clava 
and sometimes F6 brown; remainder of antenna 
yellow; dorsum of thorax with a strong purple 
and blue-green lustre; scape about 3-^1 x times as 
long as broad; frontovertex between one-sixth 
and nearly one-quarter head width; costal cell 
(Fig. 26) usually gradually tapering to its apex 
and without an apical incision before the mar- 
ginal vein (Fig. 27), rarely distinctly incised api- 
cally; lower part of outer margin of forewing 
slightly convex; apex of last tergite of gaster with 
a median incision (Fig. 49); ovipositor as in Figs 
54, 58. Male: unknown, but possibly sp. O. 

Hosts. Unknown. 

Distribution. Mexico, Costa Rica, Trinidad, 
Peru, Brazil. 

Material examined. 

Type material. Holotype °- MEXICO, Tres 
Marias Islands, Magdalena I., 19. v. 1925 (H.H. 
Kiefer) (USNM Type No 51989). Paratypes: 1$, 
Tres Marias Islands, Maria Madre I., Arroyo 
Honda, 17.V.1925 (H.H. Keifer); 1$, Magdalena 
I., 20.V.1925 (H.H. Keifer). Holotype in USNM, 
paratypes in BMNH. 

Non-type material. COSTA RICA, 15$, Gua- 
nacaste, Santa Rosa NP, various localities, vari- 
ous dates ll.v. 1985-21. ii. 1987 (Janzen & 
Gauld); 1$, Alajuela, Pehas Blancas, rainforest, 
hi. 1987, (E. Cruz); 2$, Heredia, La Selva BS, 
50m, 22.i-3.ii. 1991 (J.S. Noyes); 1$, Heredia, 
3Km S. Puerto Viejo, OTS-La Selva, 100m, 
ii-iii.1993 (P. Hanson); 59, Puntarenas, Osa 
Peninsula, Puerto Jimenez, 10m, various dates 
x.l990-ii.l992 (P. Hanson, C. Godoy); 3$, Pun- 
tarenas, RF Golfo Dulce, 3Km SW Rincon, 10m, 
x-xii.1990, viii.1991, ii.1992 (P. Hanson); TRIN- 
IDAD, 1$, I.C.T.A., on cocoa, iii.1953 (F.D. 
Bennett); 4$, St George, various localities and 
dates, vi-vii.1976 (J.S. Noyes); 1$, St George, 
St Augustine, x.1976 (F.D.Bennett); PERU, 1$, 
Madre de Dios, Tambopata Res., 12°50'S 
69°20'W, 17.x-l.xi.1983 (N.E. Stork); BRAZIL, 
1$, Bahia, Fazenda Sombra das Neves, 9.x. 1978 
(F.P. Benton). Material in BMNH, INBio, CNC. 

Comments. The Costa Rican material listed 
above differs from the holotype of insularis in the 



shape of the apex of the costal cell. In the 
holotype the costal cell is clearly incised apically 
whilst in the Cost Rican material the costal cell 
gradually tapers to its apex. However, the two 
paratypes of insularis examined exhibit a more or 
less intermediate state and therefore we are 
treating the Costa Rican material as insularis. 

Aenasius insularis is close to cirrha and paulis- 
tus (see comments under cirrha). It is closest to 
paulistus and may prove to be synonymous. For 
the present we are separating insularis and 
paulistus on the degree to which the large pilifer- 
ous punctures extend between the eyes and the 
facial impression (see key). 

Aenasius kerrichi sp. n. 

Diagnosis. Female: antenna with scape mostly 
yellow, funicle yellow and remainder dark 
brown; scape about 3 x as long as broad; sensory 
area of clava inconspicuously divided in apical 
half; frontovertex over one-third head width; 
piliferous punctures below anterior ocellus shiny; 
scrobes shallow, not sharply delimited; forewings 
infuscate in proximal two-thirds or so, apex hya- 
line; apex of venation without a naked, hyaline 
streak; lower part of outer margin of forewing 
convex; apex of last tergite of concave with a 
distinct median incision. 

Female. Length: 1.65 mm. Frontovertex dark, 
metallic blue-green with a slight purple sheen, 
especially on antennal scrobes and temples; 
scape mostly yellow but faintly margined brown 
proximally; pedicel dark brown; funicle yellow; 
clava, except sensory area, dark brown; dorsum 
of thorax blackish, weakly metallic dark blue, 
axillae a little purplish; tegulae and sides of 
thorax dark brown; forewings almost uniformly 
infuscate in slightly more than basal two-thirds, 
apex hyaline; coxae, femora and tibiae dark 
brown, the femora and tibiae apically slightly 
paler; tarsi pale yellow basally tending to amber 
apically, pretarsi dark brown; gaster dark brown 
with a slight brassy sheen. 

Head with large, conspicuous, shiny piliferous 
punctures, each separated by a sharp ridge, 
punctures in ocellar area shallower and less 
shiny, between facial impression and anterior 
ocelli these punctures shiny and about the same 
diameter as anterior ocellus, or perhaps slightly 
larger; three lines of punctures between eyes and 
facial impression; facial impression relatively 
shallow and not sharply delimited; head in side 
view slightly less than twice as long as deep, and 
fairly evenly rounded from occipital margin to 
mouth margin; scape about 3 x as long as broad; 



ENCYRTIDAE OF COSTA RICA 



131 



clava longer than funicle and pedicel together, its 
apical sensory part nearly two thirds as long as 
clava itself and divided inconspicuously by a 
hardly oblique line in apical half; ocelli forming a 
strongly obtuse angle. Relative measurements: 
HW 126; FV 49; EL 65; EW 50; MS 35; SL 46; 
SW 15; other proportions of antenna similar to 
those of insularis (Fig. 13). 

Thorax dorsally with piliferous punctures dis- 
tinct but small, each separated by about their 
own diameters; sculpture on mesoscutum regular 
imbricate-reticulate; scutellum with similar but 
slightly coarser sculpture; scutellum about one- 
sixth broader than long and with a distinct mar- 
ginal carina dorsally in apical one-fifth or so; mid 
tibial spur about as long as mid basitarsus; forew- 
ing with marginal and postmarginal veins about 
as long as stigmal; costal cell relatively broad, 
subrectangular, and abruptly incised at apex, 
with three lines of setae dorsally; submarginal 
vein with a distinct subapical hyaline break; apex 
of venation without a naked, hyaline streak, 
lower part of apical margin slightly convex. Rela- 
tive measurements (holotype): FWL 250, FWW 
126, CL 98, CW 19; HWL 170, HWW 78. 

Gaster with last tergite apically biconvex either 
side of a short median incision. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Costa Rica. 

Material examined. 

Type material. Holotype °- COSTA RICA, San 
Jose, 3200m, Cerro de la Muerte, 3.iii-7.iv.l985 
(L. Masner & H. Goulet). In CNC. 

Comments. This species can be immediately 
separated from all other species of Aenasiiis by 
the relatively wide frontovertex (more than one- 
third head width), scape about 3 x as long as 
broad and the relatively broad, subrectangular 
costal cell of the forewing. 

Aenasius pelops sp. n. 

(Figs 14, 28, 29, 57, 100) 

Diagnosis. Female: antennal scape largely yel- 
low marked with brown proximally; pedicel and 
clava dark brown; funicle mostly yellow but with 
Fl-2 brownish; scape about 3 x as long as broad; 
sensory area of clava unequally divided by a 
sigmoid line running almost its whole length; 
frontovertex between one-third and one-quarter 
head width; piliferous punctures below anterior 
ocellus slightly shiny, but not smooth; facial 
impression shallow and not sharply delimited; 



forewings infuscate proximally, apical one- 
quarter or so hyaline; apex of venation without a 
naked, hyaline streak; marginal vein about as 
long as stigmal; costal cell less than 6 x as long as 
broad, subrectangular and apically incised; lower 
part of outer margin of forewing straight; apex of 
last tergite of gaster biconvex either side of a 
median incision. 

Female. Length: 1.49-1.84 mm (holotype, 1.84 
mm). Ocellar area and area above facial impres- 
sion metallic blue-green; posterior ocellar area 
dark coppery-purple; temples and scrobes metal- 
lic green with a slight brassy sheen; piliferous 
punctures between anterior ocellus and facial 
impression not very shiny, dull; setae on fron- 
tovertex translucent brown; antenna (Fig. 14) 
with radicle dark brown, scape largely yellow but 
proximally and along margins, dark brown; 
pedicel dark brown; funicle yellow, but with Fl-2 
brownish; dorsum of thorax blackish, slightly 
shiny with a slight purple and blue sheen; setae 
dark brown; tegulae dark brown; sides of thorax 
brown; forewings (Fig. 28) brown proximally, 
but with apical one-quarter, or so, hyaline; apex 
of venation without a naked, hyaline streak; 
hindwing hyaline; all coxae dark brown; femora 
dark brown, hind femora distinctly paler api- 
cally; fore and mid tibia dark brown with apices 
yellowish; hind tibia mostly pale brown, but dark 
brown along margins; tarsi amber, pretarsi and 
mid tibial spur dark brown; gaster dark brown 
with an inconspicuous blue or purple sheen. 

Head with conspicuous, relatively shallow, 
dull piliferous punctures, those in ocellar area 
shallower and very dull; punctures between facial 
impression and anterior ocellus slightly smaller 
than ocelli; two rows of punctures extend 
between eyes and facial impression, these reach- 
ing almost half way to malar sulcus from top of 
facial impression; scrobes not sharply delimited; 
head in side view nearly 2.25 x as long as deep, 
generally evenly rounded from occipital to 
mouth margin; clava slightly longer than funicle 
and pedicel together, its apical sensory part 
about three-fifths as long as clava and unequally 
divided by a slightly sinuate line running almost 
its whole length; ocelli forming a hardly obtuse 
angle; mandible with three teeth (Fig. 100). 
Relative measurements (holotype): HW 128; FV 
36; EL 80; EW 59; MS 32; SL 51; SW 17; other 
proportions of antenna as in Fig. 14. 

Thorax with small, indistinct piliferous punc- 
tures, each separated by much more than their 
own diameters; sculpture on mesoscutum regular 
imbricate-reticulate; scutellum with similar sculp- 
ture; scutellum slightly broader than long and 



132 



J.S. NOYES ANDH. REN 



with a distinct marginal carina dorsally in apical 
one-fifth or so; mid tibial spur about as long as 
mid basitarsus; forewing with distribution of 
setae at base and proportions of venation as in 
Figs 28, 29; postmarginal and marginal veins 
about as long as stigmal; submarginal vein with a 
distinct subapical hyaline break; costal cell 
slightly less than 6 x as long as broad, subrectan- 
gular and with apex strongly incised before mar- 
ginal vein, dorsally with three lines of setae; apex 
of venation without a naked, hyaline streak, 
lower part of apical margin slightly straight. 
Relative measurements (holotype): FWL 253, 
FWW 121, CL 255, CW 45; HWL 175, HWW 73. 
Gaster with last tergite apically biconvex either 
side of a strong median incision; ovipositor as in 
Fig. 57. Relative measurements (paratype): OL 
37;MT37. 

Hosts. Unknown. 

Distribution. Costa Rica (montane areas). 

Material examined. 

Holotype $, COSTA RICA, Heredia, Vara 
Blanca, Finca Georgina, 2100m, v-vi.1990 (P. 
Hanson). Paratype: COSTA RICA, l£, Puntar- 
enas, Monteverde, 15-16. vii. 1986 (L. Masner). 
Holotype and paratype in BMNH. 

Comments. Aenasius pelops is closest to kerri- 
chi and paulistus, all three species having the 
costal cell subrectangular and conspicuously 
incised apically. It is closest to paulistus and, in 
addition to differences in venation (see key), it 
can be separated on having generally dark brown 
legs (largely amber in paulistus) and costal cell of 
the forewing less than 6 x as long as broad 
(nearly 7 x as long as broad in paulistus). This 
species can be separated from kerrichi by the 
relatively narrower frontovertex (more than one- 
third head width in kerrichi). 

Aenasius paulistus Compere 

(Figs 15,30,31,60,76) 

Aenasius paulistus Compere, 1937: 392, 401-403. 
Holotype $, Brazil, USNM, examined. 

Diagnosis. Female (Length 1.19-1.53 mm): as 
pelops but differs as follows: antennal scape (Fig. 
15) mostly yellow but marked with brown proxi- 
mally; funicle yellow; forewing (Fig. 30) with 
marginal vein not more than two-thirds length of 
stigmal (Fig. 31); costal cell about 7 x as long as 
broad; last tergite of gaster apically concave with 
a shallow median incision; ovipositor as in Fig. 
60. Male (Length 1.10-1.16 mm): forewings hya- 
line; tarsi yellow, mid tibial spur yellow; funicle 



(Fig. 76) 2-segmented, Fl transverse, clava with 
a naked ventral ridge, about 4.5 x as long as 
broad and about 2.5 x as long as scape; fron- 
tovertex a little more than one-third head width 
and with piliferous punctures distinct but partly 
obscured by shallow reticulate sculpture, facial 
impression hardly remote from eyes; distance 
from anterior ocellus to facial impression about 
0.7 x minimum width of frontovertex. 

Hosts. Recorded as a parasitoid of Pseudococ- 
cus sociabilis Hamleton (see Flanders, 1940), and 
also laboratory reared on Pseudococcus mariti- 
mus (Ehrhorn)(Compere, 1937) and recorded 
from Pseudococcus calceolariae (Maskell) ( = fra- 
gilis Brain) and Pseudococcus longispinus 
(Targioni-Tozzetti) on Hedera helix and from 
Phenacoccus sp. on Bougainvillea (Kerrich. 
1967). 

Use in biocontrol. Imported to California 
from Brazil for the control of Pseudococcus 
maritimus (Compere, 1937). The species is possi- 
bly established (Gordh, 1979). 

Distribution. USA (California, introduced), 
Costa Rica (lower altitudes), Brazil. 

Material examined. 

Type material. Holotype 9< BRAZIL, Sao 
Paulo, ex Pseudococcus maritimus, xii.1034 (H. 
Compere) (Type No USNM 51988). Paratype: 
ld\ USA, California, Riverside, ex Pseudococ- 
cus maritimus, from Sao Paulo, Brazil, reared by 
S. Flanders. Paratype in BMNH. 

Other material. COSTA RICA, 1$, Guana- 
caste Prov., Sta Rosa NP, Hacienda 2-C; 
5-26. x. 1985 (Janzen & Gauld); 10, Santa Rosa 
NP, Boq. Hum. 11-0, 24.viii-14.ix. 1985 (Janzen 
& Gauld); BRAZIL, 1$, 10", Sao Paulo, 
4.xii.l934 (H. Compere); 19, lcf , Sao Paulo, on 
Hedera helix, captured by P. gahani and P. 
longispinus, 4.xi.l934 (H. Compere). In BMNH. 

Comments. Aenasius paulistus is very close to 
pelops and they were treated as altitudinal forms 
of the same species. However, the consistent 
differences in venation and relative width of the 
costal cell (see key and under pelops) lead us to 
believe that they as distinct species. 

As pointed out by Kerrich (1967), paulistus is 
also very close to insularis Compere described 
from Mexico. The two species differ principally 
in the relative size of the facial impression. In 
paulistus the facial impression is relatively small 
and not touching the eye margins so that the 
piliferous punctures extend at least half the way 
to the malar sulcus from the top of the impres- 
sion whilst in insularis the facial impression 



ENCYRTIDAE OF COSTA RICA 



133 



almost touches the eyes and the piliferous punc- 
tures do not extend below the dorsal margin of 
the impression. The sculpture of the facial 
impression is relatively deep and rough in insu- 
laris and the mid tibial spur is amber whilst in 
paulistus the mid tibial spur is generally dark 
brown, although in the holotype it is amber- 
brown. 

Aenasius frontalis Compere 

(Figsl, 10, 11,32,33,61,87) 

Aenasius frontalis Compere, 1937: 388, 392. 
Holotype 9* Panama, USNM, not examined 

Diagnosis. Female (length: 1.40-2.22 mm) 
(Fig. 1): frontovertex between piliferous punc- 
tures dark metallic green, blue or purplish, the 
piliferous punctures very shiny green or blue and 
contrasting strongly with the areas between 
them; antenna dark brown, usually with at least 
some funicle segments yellow, apex of scape 
testaceous-yellow; dorsum of thorax with a mod- 
erately strong but variable, green, blue or purple 
sheen; tegulae dark brown; antenna (Fig. 10) 
with scape about 2 x as long as broad, sensory 
area of clava divided by an oblique, sinuate line, 
the apical part much the smaller; head in side 
view about 2.5 x as long as deep, almost flat 
above scrobes, below this strongly curved 
towards mouth; frontovertex (Fig. 87) about 
one-quarter head width, piliferous punctures 
below anterior ocellus very smooth and shiny, 
scrobes deep, sharply delimited laterally and 
dorsally; forewings (Fig. 32) completely infuscate 
but paler towards apex; naked, hyaline streak 
present at apex of venation (Fig. 33), lower part 
of outer margin of forewing weakly emarginate; 
apex of last tergite of gaster medially incised; 
ovipositor as in Fig. 61. Male: unknown. 

HOSTS. Recorded from Ferrisia virgata on cacao 
(see below, and Bennett, 1957; Kerrich, 1967) 

Distribution. USA (Texas), Mexico, Costa 
Rica, Panama, Trinidad, Peru, Brazil. 

Material examined. 

Non-type material. USA, 1$, Texas, Cameron 
Co., Sabal Palm Grove Set., 6.vii.l982 (G.A.P. 
Gibson); 2$, Texas, Cameron Co., Brownes- 
ville, viii.1983 (M. Kaulbars); MEXICO, 1$, 
20m E of Concordia, 3000ft, viii.1964 (W.R.M. 
Mason); COSTA RICA, 23$, Guanacaste 
Prov., Santa Rosa NP, various dates and locali- 
ties, 3viii. 1985-21. ii. 1987 (D. Janzen & I.D. 
Gauld); 19 , Guanacaste Prov., Guanacaste NP, 
Est. Pitilla, yellow pan trap, hi. 1990 (J.S. 



Noyes); 1$- Puntarenas Prov., Monteverde, St 
Luis Valley, 17.viii.1986 (L. Masner); 2$, Pun- 
tarenas, Manuel Antonio NP, viii.1986 (L. Mas- 
ner); 19. Puntarenas, Manuel Antonio NP, 
23-28. viii. 1988 (L. Masner); 69, Puntarenas, 
Pen. Osa, Puerto Jimenez, 10m, various dates 
ii-x.1992 (P. Hanson); 19, Heredia, La Selva 
Biol. Sta., 3Km S P. Viejo, 10°26'N 84°01'W, 
18. v. 1990 (H.A. Hespenheide); 1$, Heredia, La 
Selva BS, 50m, ii.1991 (J.S. Noyes); 29, 3Km S. 
Puerto Viejo, OTS-LaSelva, 100m, ii-iii.1993 (P. 
Hanson); 19. Limon, RB Hitoy-Cerere, 
14-19. i. 1991, 100m, 14-19. i. 1991 (J.S. Noyes); 
PANAMA, 19, Canal Zone, Barro Colorado, 
14.vi.1982 (R.B. & L.S. Kimsey); TRINIDAD, 
l9< I.C.T.A., ex Ferrisia on cocoa, v. 1952 (F.D. 
Bennett); 29- I.C.T.A., ex Ferrisia virgata on 
cocoa, 1952-53 (F.D. Bennett); 1$, St. George, 
Point Gourde, secondary forest, 8. viii. 1976, (J.S. 
Noyes), 29, St. George, St. Augustine, Malaise 
trap, 15. vii-13. viii. 1976 (J.S. Noyes); 19, St 
George, St Augustine, wasteground, 16. vi. 1976 
(J.S. Noyes); PERU, 19, Loreto, Iquitos, 
QuistoCocha, 5.ii.l984 (L. Huggert); BRAZIL, 
29 , Bahia, Mucuri, xi. 1978 (F.Benton). Material 
inBMNH, INBio. CNC 

Comments. Aenasius frontalis can be separated 
from other species of the genus by the characters 
given in the key, most notably the very character- 
istic appearance of the piliferous punctures on 
the frontovertex (see diagnosis). 

Aenasius bolowi Mercet 

(Figs 4, 6, 34, 35, 64, 71, 88, 92, 96, 102) 

Aenasius bolowi Mercet, 1947: 466-467. Lecto- 
type 9- Costa Rica, IEE, examined. 

Aenasius similis Kerrich, 1967: 196-197, 219. 
Holotype 9- Panama, USNM, examined, syn. 
n. 

Diagnosis. Female (length: 1.40-2.50 mm): 
head generally dull, metallic green; thorax black 
with a dull metallic blue or purplish sheen; 
antenna uniformly blackish; tegulae dark brown; 
antenna (Figs 4, 6) with scape about 1.6 x as 
long as broad, sensory area of clava divided by a 
slightly oblique longitudinal line (Fig. 92); head 
in side view evenly curved to top of scrobes, sides 
of facial impression slightly bulging outwards and 
then abruptly angled inwards towards mouth; 
frontovertex (Fig. 88) about one-quarter head 
width, piliferous punctures below anterior ocel- 
lus dull, scrobes deep, sharply delimited dorsally 
and laterally; forewings (Fig. 34) hyaline or dis- 
tinctly paler towards apex with hyaline streak 



134 



J.S. NOYES ANDH. REN 



present at apex of venation (Fig. 35), lower part 
of outer margin of forewing straight; apex of last 
tergite of gaster slightly concave. Male (length: 
0.88-1.78 mm): forewings hyaline (Fig. 64); tarsi 
brown, the basal segment proximally yellowish, 
mid tibial spur dark brown; funicle (Figs 71, 102) 
6-segmented, pedicel longer than Fl and about 
as wide as Fl or narrower, funicle segments very 
broad appearing oblique and each with a con- 
spicuous ventral projection, F6 and clava with 
distinct ventral oval sensory areas (Fig. 102), 
clava about 1.5 x as long as broad; frontovertex 
about one-third head width with distinct pilifer- 
ous punctures descending part way between eyes 
and facial impression (Fig. 96), but facial impres- 
sion remote from eyes; distance from anterior 
ocellus to facial impression about equal to mini- 
mum width of frontovertex. 

Hosts. Unknown. 

Distribution. Guatemala, Belize, Costa Rica, 
Panama, Grenada, Trinidad, Venezuela, Ecua- 
dor, Peru, Brazil. 

Material examined. 

Type material. Lectotype 9 °f Aenasius bolowi: 
COSTA RICA, San Mateo (Bollow) (IEE). 
Holotype 9 °f Aenasius similis Kerrich: 
PANAMA, 1$, Montilerio, v. 1924 (Fullaway) 
(Type No 71128 USNM); paratypes: GUATE- 
MALA, 1$, on banana debris (at Philadelphia, 
USA), 20.vii.1934; PANAMA, 29, Montelirio, 
v.1924 (Fullaway); 19, unrealised, vii.1914 
(D.T. Fullaway); 19, Canal Zone, Paraizo, 
20.iii.1911 (E.A. Schwarz); 19, unlocalised, on 
banana, viii.1932; 19, Barro Colorado Isl., run- 
ning on banana leaf, viii.1932 (D.T. Fullaway); 
29, Canal Zone, summit, xi.1946 (N.L.H. 
Krauss); 1$, Aquadulce, xi.1946 (N.L.H. 
Krauss); VENEZUELA, 1$, San Esteban, 
xi.1939 (P. Anduze); PERU, 1 9, reared at South 
American Parasite Lab. (Berry). Paratypes in 
BMNH, USNM. 

Non-type material. BELIZE, 29, San Ignatio, 
21.vii.1978 (P.S. Broomfield); 1$, Toledo, 25m 
NW of Punta Gorda, Salamanca, 
28.viiM.ix.1978 (P.S. Broomfield); COSTA 
RICA, 679, 380", Guanacaste Prov., Santa 
Rosa NP, various localities and dates 1985-1991 
(Gauld & Janzen); 19, Guanacaste Prov., Santa 
Rosa NP, 300m, v. 1988 (Gauld & Mitchell); lef , 
Guanacaste, 13 Km E. Filadelfia, Hda El Viejo, 
40m, v-vi.1989 (M. Garcia); 149, 8cf, Guana- 
caste Prov., Guanacaste NP, various localities, 
2-20.iii.1990 (J.S. Noyes); 1$, Guanacaste 
Prov., Palo Verde, dry forest, 4-12. vi. 1988 (B.V. 
Brown); 19, Guanacaste Prov., NW Volcan 



Orosi, Cerro el Hacha, 1988; 19, San Jose, San 
Pedro, Tigra Cacao, iii— iv. 1990 (P. Hanson); 19, 
San Jose, Ciudad Colon, iii— iv. 1990 (P. Hanson); 
19, 4cf, San Jose, Ciudad Colon, Hda El 
Rodeo, 16. ii. 1991 (Hym. Parataxonomists); 59, 
18d\ Heredia Prov., La Selva Biol. Sta., 3 km S. 
Pto Viejo, 10°26'N 84°01'W, on leaves of Alchor- 
nea costaricensis Pax & Hoffm., various dates 
23.iii-7.iv.1987 (H.A. Hespenheide); 59, 4<3\ 
Heredia Prov., La Selva Biol. Sta., 3 km S. Pto 
Viejo, 10°26'N 84°01'W, 7.iv.l987 (H.A. 
Hespenheide); 19, lef, Puntarenas, Manuel 
Antonio NP, 23-28. viii. 1986 (L. Masner); 19, 
Puntarenas, Osa Peninsula, 80 mi. ( = 128 km) 
SW Rincon, 08°42'N 83°29'W, iii. 1987 (H.A. 
Hespenheide); 19, Puntarenas, Osa Peninsula, 
Puerto Jimenez, 10m, x-xi.1990 (P. Hanson); 
19, 2d", Puntarenas, Golfito, 20.iii.1990 (J.S. 
Noyes); lef, Cartago, Turrialba, CATIE, 700m, 
14-15. iii. 1980 (J. S. Noyes); 19, Cartago, San 
Ramon, Tres Rios, 1500m, on Conostegia 
xalepeh, 2. vi. 1990 (P. Hanson); 1$, Limon, 7Km 
SW Bribri, ix-xi.1989 (P. Hanson); 1$, Limon, 
RB Hitoy-Cerere, 100m, 14-19.U991 (J.S. 
Noyes); PANAMA, lef. Canal Zone, on banana 
trees, iv.1924 (D.T. Fullaway); 1$, Canal Zone, 
Barro Colorado I., 14. vi. 1982 (R.B. & L.S. 
Kimsey); 19, Santiago de Veraquas, 27.vii.1976 
(F.D. Bennett); GRENADA, lef, Granville 
(windward side), 36 (H.H. Smith); TRINIDAD, 
19, Caroni, Caroni Swamp, 1 .vii. 1976 (J.S. 
Noyes); ECUADOR, 29 , Pichinchas, Tinlandia, 
800m, 8.ii.l983 (L. Huggert); BRAZIL, 29, 
Amazonas, Manaus, Campus Univ., 29. vi. 1982 
(J.A. Rafael). Material in BMNH, USNM, 
INBio. 

Comments. Aenasius bolowi is very close to 
hyettus (Figs 7, 36) and may be synonymous. We 
are only able to separate the two species on the 
difference in the shape of the scape as noted by 
Kerrich (1967), which in some cases is very small 
and may be related to body size. In general 
specimens from St Vincent and Grenada (hyet- 
tus) are smaller and tend to have the distal part of 
the scape more evenly tapering (Fig. 7) whilst 
those from elsewhere (bolowi) are larger and 
tend to have the distal part of the scape slightly 
bulging outwards distad of the pedicel insertion 
(Fig. 6). For the present we prefer to treat the 
two species as distinct until we are able to 
examine larger specimens from St Vincent to 
assess the range of variation more accurately. 

Aenasius bolowi is superficially very similar to 
caeruleus but can be separated by having the 
apical part of the scape not very strongly bulging 
outwards, the scrobal impression dorsally sharply 



ENCYRTIDAE OF COSTA RICA 



135 



margined, different sculpture of the antennal 
scrobes and interantennal prominence (compare 
Figs 88 and 89), piliferous punctures below the 
anterior ocellus relatively dull, and the division 
of the sensory part of the clava more or less 
longitudinal and straight (Fig. 92). A. caeruleus 
has the apical part of the scape distinctly bulging 
outwards, the scrobal impression not sharply 
margined, the piliferous punctures of the fron- 
tovertex very shiny below the anterior ocellus 
and the division of the sensory part of the clava 
distinctly transverse and sigmoidal (Fig. 93). 
There are also further differences between these 
two species in the shape of the apex of the outer 
part of the base of the second valvifers and shape 
of the hypopygium. 

Aenasius caeruleus Brues 

(Figs 3, 38, 39, 89, 93) 

Aenasius caeruleus Brues, 1910: 84-85. Holotype 

9, Mexico, AMNH, examined. 
Aenasius personatus Kerrich, 1967: 198, 199, 

221. Holotype 9, USA, USNM, examined. 

syn. n. 

Diagnosis. Female (length: 1.01-2.30 mm): 
head metallic violet to green, ridges between 
piliferous punctures occasionally purplish; 
antenna more or less uniformly dark brown; 
dorsum of thorax with a moderate metallic 
green, blue, purple or violet sheen, occasionally 
hardly metallic; tegulae dark brown, normally 
with a blue or purple sheen; antenna (Fig. 3) with 
scape widest beyond middle, 1.4-1.9 x as long as 
broad, sensory area of clava divided obliquely by 
a sinuate line (Fig. 93); head in side view about 2 
x as long as deep, evenly curved above scrobes, 
straight below this but sometimes distinctly 
angled towards mouth below eyes; frontovertex 
(Fig. 89) about one-third to less than one-quarter 
head width, piliferous punctures below anterior 
ocellus shiny, sometimes extremely so; scrobes 
moderately deep, not sharply delimited dorsally; 
eyes with conspicuous dark hairs; forewings (Fig. 
38) with a naked, hyaline streak at apex of 
venation (Fig. 39), costal cell with at least three 
lines of setae dorsally, lower part of outer margin 
of forewing straight; apex of last tergite of gaster 
hardly concave. Male: Probably very similar to 
that described for maplei Compere and may be 
that described here as sp. M. 

Variation. Notable variation mentioned under 
the diagnosis includes the variation in the relative 
width of the frontovertex and scape and colora- 
tion of the head and thorax. There is also some 



significant variation in the relative length of the 
sensory area of the clava, in some specimens it is 
only about twice as long as the distance separat- 
ing it from the base of the clava, whilst in other it 
is about three times as long. The sculpture of the 
mesoscutum also varies from quite smooth to 
finely reticulate. 

Hosts. Recorded below and by Kerrich (1967) 
as a parasitoid of Ferrisia virgata (Cockerell) on 
cacao and Gliricidia, and from Ferrisia sp. on 
Gliricidia. 

Distribution. USA (Texas, Florida), Mexico, 
Belize, Costa Rica, Panama, Puerto Rico, Trin- 
idad, Venezuela, Ecuador, Peru, Brazil, Uru- 
guay. 

Material examined. 

Type material. Holotype 9 of caeruleus. Type 
No. A.M.N.H., Am. Mus. Nat. Hist. Dept. 
Inver. Zool. No 21120, 411 Petrun, Aenasius 
caeruleus Brues Type (according to Brues, the 
specimen was collected in Vera Cruz, MEXICO 
by A. Petrunkevitch) (AMNH). Holotype 9 of 
personatus. USA, Florida, Hialeah, on Hibiscus 
tiliaceus 24.viii.1953 (O.D. Link) (USNM). 
Paratypes: USA, 29. Florida. Hialeah, on Hibsi- 
cus tiliaceus, 24.viii.1953 (O.D. Link); TRIN- 
IDAD, 39. I.C.T.A., ex Ferrisia virgata on 
cocoa, iii.1952 (F.D. Bennett); 19, I.C.T.A., ex 
Ferrisia virgata on cocoa, 1952-1953 (F.D. Ben- 
nett); 19. I.C.T.A., ex Ferrisia on Gliricidia, 
ix. 1953 (F.D. Bennett); 19. San Juan, ex Ferrisia 
on Gliricidia, xi. 1953 (F.D. Bennett). Paratypes 
in BMNH. 

Non-type material. USA, 29. Texas, Brewster 
Co., Big Bend NP, 23-28.vi.1982 (G.A.P. Gib- 
son); MEXICO, 19. Nuevo Leon, San Pedro, 
Garza Garcia, 20.ix. 1983 (M. A. Rodriguez); 1$, 
Vera Cruz, 33Km NE Catemaco, vii.1983 (M. 
Kaulbars); 19. Chicana, Ruins 6m E Xpujil, 
vii.1983 (R. Anderson); BELIZE, 1$, Han- 
lover, 10.vii.1978 (P.S. Broomfield); COSTA 
RICA, 49, Guanacaste Province, Santa Rosa 
NP, Hacienda 1-0, various dates viii. 1985-x. 1986 
(D. Janzen & I.D. Gauld); 19, Alajuela, ex 
mealybug on M. aesculifolia, 6. v. 1981 (CIE 
13352); 19, Alajuela, Chiles de Aguas, Zarcas 
cafe, xii.1989 (R. Cespedes); 1$, Heredia, La 
Selva Biol. Sta., 3 Km S P. Viejo, 10°26'N 
84°01'W, 17. iv. 1988 (H.A. Hespenheide); 19, 
Heredia, 3Km S. Puerto Viejo, OTS-La Selva, 
100m, ii— iii.1993 (P. Hanson); 29, Cartago, La 
Cangreja, 1950m, vii.1991 (P. Hanson, C. 
Godoy); 29, Puntarenas Prov., Manuel Anto- 
nio, 23-28. viii. 1988 (L. Masner); 39, Puntar- 
enas, San Vito, Est. Las Alturas, 1500m, 



136 



J.S. NOYES AND H. REN 



i-ii.1992 (P. Hanson); PANAMA, 1$, Canal 
Zone, Paraiso, 6.ii. 191 1 (E.A. Schwarz); 
PUERTO RICO, 1$, Mun Dorado, beach 2Km 
W. Pt. Salinas, 25.V.1982 (S.L. Heydon); TRIN- 
IDAD, 1$, I.C.T.A., ex Ferrisia sp. on cocoa, 
v.1952 (F.D. Bennett) (paratype of Aenasius 
regularis Kerrich, misidentification); lcf, 
I.C.T.A., ex Ferrisia on Gliricidia, ix.1953 (F.D. 
Bennett) [det G.J. Kerrich, 1967; probable misi- 
dentification of bolowi]; 1$, I.C.T.A., ex Ferri- 
sia virgata on Gliricidia, vi.1955 (F.D. Bennett); 
l^, Caroni, Gran Couva, wasteground, 
16.vii.1976 (J.S. Noyes); 1$, St. Patrick, Coora, 
coffee plantation and rainforest, 14.viii. 1976 
(J.S. Noyes); 1$, St. George, St. Augustine, 
wasteground, 18. vi. 1976 (J.S. Noyes); 1$, St. 
George, St. Augustine, Malaise trap, 
15.vii-13.viii.1976 (J.S. Noyes); 1$, Curepe, Sta 
Margarita Circular Rd., 21-23.xi.1977 (W.R.M. 
Mason); ECUADOR, 1$, Rio Palenque, forest, 
4.H.1983 (L. Huggert); 2$, Napo, 10km N.E. 
Tema, Rio Hollin, 400m, 19.ii.1983 (L. Masner); 
1$, Napo, Misahualli, 20.ii.1983 (L. Huggert); 
2$, Napo, Sacha, 7.iii.l983 (L. Huggert); 
PERU, 1$, Cuzco, Quilambamba, 
24-26.xii.1983 (L. Huggert); 3$, Madre de Dios, 
Tambopata Res., 12°50' 69°20'W, 17.x-l.xi.1983 
(N.E. Stork); 1$, Huanuca, Tingo Maria, 
26.U984 (L. Huggert); BRAZIL, 2$, Santa 
Catarina, Nova Teutonia,xi. 1949, 19.xii.1951 (F. 
Plaumann). Material in BMNH, INBio, CNC. 

Comments. Included here are specimens which 
vary considerably in the relative width of the 
scape and frontovertex and in the length of the 
sensory area of clava and general coloration of 
the body. Initially, we considered that more than 
one species might be represented, but we now 
believe that only one variable species is present. 
The differences between caeruleus and per- 
sonatus fall within the limits of this variation and 
therefore we treat the two names as synonymous. 
Aenasius maplei Compere is also very close to 
caeruleus and may be synonymous, but we hesi- 
tate to formally propose this synonymy until 
further North American material can be exam- 
ined. 

The close relationship of caeruleus with maplei 
leads us to assume that the male of caeruleus is 
probably very similar to that of maplei (see 
Compere, 1937). This suggests that Aenasius sp. 
M may be the male of caeruleus and not that 
identified as personatus by Kerrich (BMNH col- 
lection) which we are sure is the male of hyettus. 

Aenasius caeruleus can be confused easily with 
hyettus (see comments under hyettus) and regu- 
laris Kerrich. Females of caeruleus can be sepa- 



rated most conveniently from those of regularis 
by the less strongly curved stigmal vein (compare 
Figs 37 and 39). 

Aenasius brasiliensis (Mercet) 
(Figs 16, 40, 41, 65, 69, 90) 

IBlepyrus tachigaliae Brues, 1921: 229-230. 

Holotype $, Guyana, ?lost. 
Chalcaspis brasiliensis Mercet, 1926: 46-48. 

Holotype $, Brazil, IEE, examined. 
Aenasius cariocus Compere, 1937: 390, 399, 404. 

Holotype 9- Brazil, USNM, examined, syn. n. 
Aenasius colombiensis Compere, 1937: 403-404. 

Holotype $, USNM, not examined, syn. n. 

Synonymised with cariocus by Kerrich, 1967. 
Aenasius brasiliensis (Mercet); Compere, 1937: 

390, 398. 
Aenasius theobromae Kerrich, 1953: 796-797. 

Holotype $, Trinidad, BMNH, examined. 

syn. n. Synonymised with cariocus by Kerrich, 

1967. 

Diagnosis. Female (length: 1.21-1.90 mm): 
head generally metallic green or blue green, 
ridges between piliferous punctures purplish, 
ocellar area duller; antenna with scape yellow, 
variously marked with brown proximally and 
along dorsal and ventral margins; pedicel and 
clava dark brown; funicle segments yellow some- 
times marked extensively with brown; dorsum of 
thorax relatively dull, but with a slight greenish 
or purple sheen; tegulae dark brown; antenna 
(Fig. 46) with scape normally about 3-3.5 x as 
long as broad, sensory area of clava divided 
obliquely into two almost equal halves; head in 
side view about 2 x as long as deep, almost flat 
above scrobes, below this more strongly, curved 
towards mouth; frontovertex (Fig. 90) between 
one-quarter and one-third head width, piliferous 
punctures between anterior ocellus and facial 
impression shiny, scrobes moderately deep, not 
sharply delimited; forewings (Fig. 40) with apical 
half hyaline, or at least conspicuously paler than 
basal half; naked, hyaline streak at apex of 
venation present (Fig. 41); lower part of outer 
margin of forewing straight; apex of last tergite 
of gaster hardly concave and not medially 
incised; ovipositor about two-thirds as long as 
mid tibia. Male (length: 0.65-01.27 mm): forew- 
ings normally with base infuscate (Fig. 65), rarely 
completely hyaline; tarsi yellow, mid tibial spur 
dark brown; scape (Fig. 69) about 2.5 x as long 
as broad and widest in middle, funicle 
6-segmented, pedicel longer than Fl and about 
as wide, funicle segments disc-like and without a 
conspicuous ventral projection, F6 with an oval 



ENCYRTIDAE OF COSTA RICA 



137 



sensory area, clava about twice as long as broad; 
frontovertex about one-third head width with 
distinct piliferous punctures which do not 
descend between eyes and facial impression, but 
facial impression remote from eyes; distance 
from anterior ocellus to facial impression about 
0.65-0.8 x minimum width of frontovertex. 

Hosts. Recorded from Pseudococcus sp. in Bra- 
zil and Colombia (Compere, 1937; Kerrich, 
1967) and from Dysmicoccus brevipes (Cocker- 
ell) in Trinidad (Bennett, 1957; Kerrich, 1953, 
1967). 

Use in biocontrol. The species was apparently 
introduced into Hawaii in 1935 for the control of 
Dysmicoccus brevipes (Cockerell) (Swezey et ai, 
1939, as colombiensis), but no further informa- 
tion is available. 

Distribution. Mexico, Costa Rica, Panama, 
Trinidad, Colombia, Ecuador, Peru, Brazil, 
Bolivia. 

Material examined. 

Type material. Holotype 9 of Chalcaspis brasil- 
iensis: BRAZIL, Corumba, Matt. Grosso (IEE). 
Holotype 9 of Aenasius cariocus: BRAZIL, 
Campinas, 26. xi. 1934 (H. Compere) (Type No 
51987 USNM) (antenna and wings on slide, but 
slide missing). Paratypes: A. colombiensis, 
COLOMBIA, 3°, 2o\ Barbosa, x.1935 (E.G. 
Salas); A. theobromae, TRINIDAD, 1$, 2d\ 
Maracas, ex Pseudococcus brevipes on cacao 
pod, x.1949 (T.W. Kirkpatrick). Unless other- 
wise stated material in BMNH. 

Non-type material. MEXICO, 19, Vera Cruz, 
20 Km NE Tianchinol, vi.1983 (M. Kaulbars); 
19, Campos, 10 Km W Xpulil, Chincanna, 
300m, viii.1983 (M. Kaulbars); COSTA RICA, 
99, Guanacaste Prov., Santa Rosa NP, various 
localities and dates, vii.1985-ix.1991 (Janzen & 
Gauld); 19, 2d\ Guanacaste NP, near HQ, and 
7Km E. HQ, 2-10.iii.1990, (J.S. Noyes); 79, 
lo\ Alajuela, Rio Pefias Blancas, 10°19'N 
84°43'W, 800m, 2517, ex Cataenococcus sp. 
[det. D.J. Williams] in stems of Cecropia insignis 
inhabited by Azteca xanthochroa, 10-13. v. 1989 
(J. Longino); 19, Alajuela, Rio Pefias Blancas, 
18.vii.1986 (L. Masner); 49, Puntarenas, 4Km 
E. Palmar Norte, 8°58'N 83°25'W, 40m, 2665, 
ex Pseudococcus near neobrevipes [det. D.J. 
Williams] in domatium Cordia aliodera, nest of 
Crematogaster curvispinosa, 26.iii.1990 (J. 
Longino); 19, Puntarenas, Golfo Dulce, 3Km 
DW Rincon, 10m, ii-v.1989 (P. Hanson); 29, 
San Jose, Ciudad Colon, Hda El Rodeo, 
18. ii. 1991 (Hym. Parataxonomists); 19. 



Heredia, Braulio Carillo NP (HQ), 250-500m. 
10.vi.1985 (L. Masner, H. Goulet); 10$, 
Heredia, La Selva Biol. Sta., 3km S Porto Viejo, 
10°26'N 84°01W', v.l990-vii.l991 (H.A. 
Hespenheide); 19, Heredia, La Selva BS, 50m, 
22.i-3.ii. 1991 (J.S. Noyes); 39, Heredia, 3 Km 
S. Puerto Viejo, OTS-LaSelva, 100m, ii-iii.1993 
(P. Hanson); 79, Limon, RB Hitoy-Cerere, 
100m, 14-19.i.l991 (J.S. Noyes); TRINIDAD, 
19, Chatham, on grass, 1913 (F.W. Urich); 19, 
unlocalised, vial No 1 (F.D. Bennett); 39. Cara- 
cas Valley, ex [Dysmicoccus] brevipes on cocoa, 
No 18, iii.1953 (F.D. Bennett); 19, Curepe, Sta 
Margarita Circ. Rd., 26.ix-26.x.l974 (F.D. Ben- 
nett); 19, St George, El Tucuche (West Slope), 
26.vi.1976 (J.S. Noyes); 19, St George, St. 
Augustine, Malaise trap, 15. vii— 13. viii. 1976 (J.S. 
Noyes); 39, Arima Valley, 5th mile, edge of 
rainforest, 3. vii. 1976 (J.S. Noyes); COLOM- 
BIA, 29, Bucaramanga, ix. 1935 (E.G. Salas); 
19, Barbosa, x.1936 (E.G. Salas); 39, Vaupes, 
River Vaupes, ex coccid, x— xii. 1952 (D.J. Tay- 
lor); ECUADOR, lcf, Pichincha, Tinlandia, 
800m, 8.H.1983 (L. Huggert); 19, 2o\ Napo, 
Tena, 16. ii. 1983 (L. Huggert); 19, Napo, Mis- 
ahualli, 20.ii.1983 (L. Huggert); 29, 2d\ Napo, 
Sacha, 7.iii.l983 (L. Huggert); PERU, 889, 
15d\ Madre de Dios, Rio Tambopata Res., 
12°50S 69°2()W, 1-13. xi. 1983 (N.E. Stork); 79, 
8cf, Madre de Dios, Pto Maldonado, 3. i. 1984 
(L. Huggert); 49, 3Q\ Junin, Satipo, 
18-24.L1984 (L. Huggert); 49, Loreto, Iquitos, 
Quisto Cocha, 5.ii.l984 (L. Huggert); 29, 
Loreto, Iquitos, NE Rio Nanay, 8.ii.l984 (L. 
Huggert); 39, lO\ Loreto, Iquitos, Granja 
Unap, 9.ii.l984 (L. Huggert); 39, 10", Loreto, 
Iquitos, Barillal, 10. ii. 1984 (L. Huggert); BRA- 
ZIL, 19, Sao Paulo, ex Phenacoccus sp. on 
Boiugainvillia, 28.xii.1934 (E. Hambleton); 59, 
Sao Paulo, Guaraja, ex Pseudococcus sp 16, 
vii. 1935 (E. Hambleton); Id", Sao Paulo, ex 
Pseudococcus sp 15, viii. 1935 (E. Hambleton); 
19, Sao Paulo, Teodora Sampaio, xii. 1977 (M. 
alvarenga); 59, Santa Catarina, Nova Teutonia, 
various dates, ix.1943-x.1949 (F. Plaumann); 
39, Campinas, No 12 , ex Pseudococcus, ii. 1936 
(E. Hambleton); 19, Para, Oriximina, Alcoa 
Hine Racao, Rio Trombetas, 7-25. x. 1982 
(Rafael, Binda & Vidal); 19, Mato Grosso, 
Sinop., 19. ii. 1956 (O. Roppa); 19, Bahia, Mer- 
curi, xi.1978 (F.P. Benton); 19, Bahia, Buer- 
arema, Faz Casnue e Damaio, 2.x. 1980 (F.P. 
Benton); 29, Bahia, Itabuna, iv.1983 (F.P. Ben- 
ton); 39, Bahia, Itabuna, CEPEC, ex mealybug 
in Azteca nest in Cecropia stem, ix.1986 (F.P. 
Benton); 19, Minas Gerais, Aguas Vermelhas, 
xii. 1983 (M. Alvarenga); 19, Rondonia, Vii- 



138 



J.S. NOYES ANDH. REN 



hena, xi.1973 (M. Alvarenga); 1$, Amazonas, 
Manuas, CDC trap, 15.i-25.ii. 1981 (J. Arias); 
1$, Amazonas, Sao Gabriel de Cachoeria, 
20-29. iv. 1982 (J. Arias); 5$, Amazonas, 
Manaus, Univ. campus, Malaise trap, vi.1982 
(J. A. Rafael); BOLIVIA, 1$, Yungas, 15 Km 
NE Caranavi, 920m, 26. i. 1973 (J. Helava). Mate- 
rial in BMNH, CNC, INBio, USNM, TAMU. 

Comments. We have examined the respective 
holotypes of C. brasiliensis and A. cariocus and 
conclude that the two belong to the same species. 
The apparent difference in the relative width of 
the scape noted by Kerrich (1967) is due to the 
lamina of the scape slightly curling inwards on 
the holotype of brasiliensis which gives it the 
appearance of being narrower. 

A. brasiliensis is similar to, and probably 
closely related to, longiscapus Compere from 
which it can be reliably separated using the 
characters given in the key to species. In addition 
to the these characters the two can be separated 
on the relative length of the ovipositor. In brasil- 
iensis the ovipositor is only about 2/3 the length 
of the mid tibia whereas in longiscapus it clearly 
longer than the mid tibia. 

Aenasius mitchellae sp. n. 

(Figs 17, 42, 43, 59) 

Diagnosis. Female: antenna with scape mostly 
blackish, funicle with yellow and brown seg- 
ments; club dark brown; scape about 4 x as long 
as broad and longer than maximum eye width; 
sensory area of clava inconspicuously divided on 
inner side in apical half by a slightly oblique line; 
frontovertex slightly less than one-third head 
width; piliferous punctures below anterior ocel- 
lus relatively dull; scrobes shallow, not sharply 
delimited; forewings infuscate, basal area dark- 
est; apex of venation with a naked, hyaline 
streak; lower part of outer margin of forewing 
almost imperceptibly emarginate; apex of last 
tergite of concave with a distinct median incision. 

Female. Length: 2.09-2.35 mm (holotype 2.35 
mm). Frontovertex in ocellar area dark green, 
almost black and hardly shiny, below this gradu- 
ally becoming more metallic and blue-green; 
scrobes metallic green with brassy reflections and 
with some coppery reflection dorsally, interan- 
tennal prominence margined violet dorsally; 
genae with some purple reflections; scape mostly 
blackish, slightly metallic blue and purple, apex 
yellow; pedicel, Fl-3 and clava blackish, slightly 
metallic; F4-6 yellow; dorsum of thorax with a 
strong metallic blue-green lustre mixed with cop- 



pery and purple; tegulae and sides of thorax dark 
brown, almost black; forewings strongly infus- 
cate, weakest towards wing apex; coxae, fore 
femora and fore tibiae almost black; mid femur 
dark brown in basal half, apex amber; hind 
femur similar but darker; mid tibia amber; dor- 
sally dark brown; hind tibia similar but also 
margined brown ventrally; mid tibial spur dark 
brown; tarsi amber, pretarsi dark brown; gaster 
dark brown with a slight brassy, blue and purple 
lustre. 

Head with conspicuous, piliferous punctures 
which are mostly dull, but becoming more shiny 
towards facial impression, those immediately 
above facial impression quite smooth; punctures 
about the same diameter as anterior ocellus, or 
perhaps slightly larger; three lines of punctures 
between eyes and facial impression; facial 
impression shallow and not sharply delimited; 
head in side view about 2.25 x as long as deep, 
and fairly evenly rounded from occipital margin 
to mouth margin; scape (Fig. 17) about 4 x as 
long as broad; clava about as long as funicle and 
pedicel together, its apical sensory part slightly 
more than two thirds as long as clava and divided 
on inner side in its apical half by a hardly oblique 
line; ocelli forming a very slightly obtuse angle. 
Relative measurements (holotype): HW 140; FV 
40; EL 81; EW 61; MS 40; SL 75; SW 18. 

Thorax dorsally with piliferous punctures 
indistinct, each separated by much more than 
their own diameters; sculpture on mesoscutum 
very fine but regular transversely elongate, 
imbricate-reticulate; scutellum with similar but 
polygonal, almost reticulate sculpture; scutellum 
slightly broader than long and with a distinct 
marginal carina dorsally in apical one-fifth or so; 
mid tibial spur about as long as mid basitarsus; 
forewing (Fig. 42) with stigmal vein nearly twice 
as long as marginal and about 1.5 x as long as 
postmarginal (Fig. 43); costal cell gradually 
tapering distally, not abruptly incised apically 
and with two lines of dorsal setae; submarginal 
vein with a distinct subapical hyaline break; apex 
of venation with a naked, hyaline streak, lower 
part of apical margin at least slightly emarginate. 
Relative measurements (holotype): FWL 320, 
FWW 140, CL 123, CW 19; HWL 213, HWW 83. 

Gaster with last tergite slightly concave and 
with a median incision; ovipositor (Fig. 59) 
nearly one quarter longer than mid tibia. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Costa Rica. 



ENCYRTIDAE OF COSTA RICA 



139 



Material examined. 

Type material. Holotype 9, COSTA RICA, 
Heredia, Vara Blanca, vii-viii.1990 (P. Hanson). 
Paratypes: COSTA RICA, 1 $, San Jose, 
Zurqui de Moravia, 1600m, ii. 1989 (P. Hanson); 
29 , San Jose, Cerro de la Muerte, 19Km S, 3Km 
W. Empalme, 2600m, iv-vii.1992, iv-v.1993 
(P.Hanson, C. Godoy). Holotype in BMNH, 
paratypes in BMNH, INBio. 

Comments. A. mitchellae is close to longiscapus 
Compere, both having a hyaline streak at the 
apex of the venation; scape more than 3 x as 
long as broad; outer margin of forewing emargin- 
ate; and frontovertex more than one-quarter 
head width. It differs from longiscapus Compere 
in the sculpture of the facial impression being 
very nearly smooth whereas in longiscapus it is 
very rough; the postmarginal vein in longiscapus 
is very nearly as long as the stigmal (clearly 
shorter in mitchellae); the scape in longiscapus is 
3-3.5 x as long as broad and generally yellow 
(blackish and more than 4 x as long as broad in 
mitchellae). Further to this the line dividing the 
sensory area of the clava in longiscapus is in the 
outer half (inner half in mitchellae). 

Aenasius longiscapus Compere 

(Figs 18, 19,44,45,91) 

Aenasius longiscapus Compere, 1937: 388-391, 
398-399. Holotype 9, USNM, examined. 

Aenasius pacificus Compere, 1937: 388-391, 
399-400. Holotype 9, USNM, examined, syn. 
n. 

Aenasius vadosus Kerrich, 1967: 214. Holotype 
9, Puerto Rico, BMNH, examined, syn. n. 

Aenasius acuminatus Kerrich, 1967: 215. Holo- 
type 9- Trinidad, BMNH, examined, syn. n. 

Diagnosis. Female (length: 1.19-2.09 mm): 
head with ocellar area slightly lustrous purple- 
brown, below this metallic green, occasionally 
slightly coppery or brassy; antenna mostly yellow 
usually with proximal two-fifths of scape nar- 
rowly margined dark brown ventrally, pedicel 
dark brown and clava completely grey-brown or 
brown (Fig. 18) sometimes proximal segment 
yellowish (Fig. 19); dorsum of thorax blackish 
with a dull green, blue or purple sheen; tegulae 
blackish; antenna (Figs 18,19) with scape about 
3-5 x as long as broad, sensory area of clava 
divided by an oblique, slightly sinuate line in 
apical half or so; head in side view about 2.5 x as 
long as deep, gradually and evenly curved above 
scrobes, below top of scrobes angled towards 
mouth at about 45-60°; frontovertex (Fig. 91) 



between one-third and one-quarter head width, 
piliferous punctures below anterior ocellus dull, 
but above scrobes and between eyes and scrobes 
moderately shiny; scrobal area small and rela- 
tively shallow, from mouth margin only about 
one-quarter head length, sharply delimited later- 
ally; forewings (Fig. 44) with hyaline streak 
present at apex of venation (Fig. 45), lower part 
of outer margin of forewing clearly emarginate; 
apex of last tergite of gaster hardly concave, but 
with a distinct median incision; ovipositor clearly 
longer than mid tibia. Male: unknown with cer- 
tainty, but a slide mounted specimen (see below 
under material examined) identified by Compere 
as this species is identical to males of phenacocci 
and advena (see diagnosis for advena). 

Variation. A variable species, both in the 
shape and coloration of the scape, coloration of 
the clava (Figs 18, 19), head shape, relative 
length and shape of the hypopygium and length 
of the ovipositor. Variation in coloration and 
relative width of the scape has been discussed 
above. Variation in the shape of the hypopygium 
is related to the relative length of the ovipositor. 
In specimens where the ovipositor is about 1.5 x 
as long as the mid tibia the hypopygium is 
significantly more elongate than in specimens 
where the ovipositor is only about 1.25 x as long 
as the mid tibia. The head in facial view varies 
from being dorsally quite rounded to virtually 
straight with the dorso-lateral eye margins being 
relatively angular. 

Hosts. Recorded below from Ferrisia virgata 
(Cockerell), Ferrisia sp, on Gliricidia, and from 
Dysmicoccus brevipes (Cockerell) on cacao (see 
also Kerrich, 1967). 

Distribution. Mexico, Costa Rica, Puerto 
Rico, Puerto Rico, Trinidad, Peru, Ecuador, 
Brazil. 

Material examined. 

Type material. Holotype 9 of longiscapus: BRA- 
ZIL, Sao Paulo, Campinas, 26. xi. 1934 (H. Com- 
pere) (USNM Type No 51991). Holotype 9 of 
pacificus: MEXICO, Tres Marias Islands, Maria 
Madre Arroyo Hondo, 17.V.1925 (H.H. Kiefer) 
(USNM Type No 51993). Holotype 9 of acumi- 
natus: TRINIDAD, Maracas, ex Dysmicoccus 
brevipes (Ckll.) on cacao, v. 1953 (F.D. Bennett) 
(BMNH). Holotype 9 of vadosus: PUERTO 
RICO, Mayaguez, on coffee, xi.1959 (F.J. [sic] 
Bennett) (BMNH). 

Other material. COSTA RICA, 59, Guana- 
caste, Santa Rosa NP, Hacienda-3-0, various 
dates 22.vi. 1985-8. x. 1986 (Janzen & Gauld) ; 1 9 , 



140 



J.S. NOYES AND H. REN 



Guanacaste, Volcan Orosi, Cerro el Hacha, 
300m, 1988; 1$, Guanacaste, 13 Km E. Fil- 
adelfia, Hda El Viejo, 40m, v-vi.1989 (M. Gar- 
cia); 1$, Heredia, La Selva Biol. Sta., 50m, 
ii.1991 (J.S. Noyes); 1$, Heredia, La Selva Biol. 
Sta., 3Km S Pto, Viejo, 10°26'N 84°01'W, 
14. vi. 1991 (H.A. Hespenheide); 1$, Heredia, 3 
Km S. Puerto Viejo, OTS-La Selva, 100m, 
ix.1992 (P. Hanson); 1$, 3Km S. Puerto Viejo, 
OTS-La Selva, 100m, canopy fog, Carapa sp., 
FOT/02, 15. hi. 1993; 2$, Barra Honda, 150m, 
v.1988 (I.D. Gauld); 2?, Puntarenas, Manuel 
Antonio NP, 24.viii.986 (L. Masner); 1$, Pun- 
tarenas, Pen. Osa, Puerto Jimenez, 10m, ii.1992 
(P. Hanson); 1$, Limon, P.I. Amistad, San Vito 
C. Brus, Finca Catrosa, iv-x.1989 (M. Ramirez); 
TRINIDAD, 1$, I.C.T.A., ex F. virgata on 
cocoa, vi.1953 (F.D. Bennett); 1$, I.C.T.A., ex 
Ferrisia on Gliricidia, 2.x. 1953 (F.D. Bennett); 
1$, I.C.T.A., ex Ferrisia on cocoa, xii.1953 
(F.D. Bennett); 1$, I.C.T.A. ex Ferrisia sp. on 
Gliricidia, iv.1954 (F.D. Bennett); 1$, Waller- 
field, on Piper, xi.1958 (F.D. Bennett); ECUA- 
DOR, 1$, Pichincha, Tinlandia, 800m, 2.ii.l983 
(Masner & Sharkey); PERU, 1$, Madre de 
Dios, Pto Maldonado, 3. i. 1984 (L. Huggert); 
BRAZIL, lcf, Sao Paulo, captured near mealy- 
bug on grass, 6.xi.l937 (H. Compere) (identified 
as longiscapus by Compere, but probably male of 
advena); 1$, Bahia, ex Ferrisiana virgata on 
coconut, xi.1964 (F.D. Bennett); l£, Bahia, 
Mucuri, xi,1978 (F.D. Bennett). Material in 
BMNH, CNC. 

Comments. The relative widths of the scape of 
pacificus and acuminatus given by Kerrich (1967) 
are incorrect. We have measured the scape in the 
holotypes of both and they are almost identical 
being 4 x and 4.3 x as long as broad respec- 
tively. Kerrich also noted several other differ- 
ences between these species, longiscapus and 
vadosus. However, the material we have been 
able to examine indicates that these differences 
are unreliable and probably reflect infraspecific 
variation. We are therefore treating all four 
names as synonyms. 



Aenasius sp. A 

(Fig. 72) 

Diagnosis. Length 1.02-1.59 mm. Forewings 
with base infuscate; tarsi yellow, mid tibial spur 
dark brown; scape widest below middle; funicle 
(Fig. 72) 6-segmented, pedicel longer and wider 
than Fl and F2 together, these two segments 
subquadrate; funicle segments disc-like and with- 
out a conspicuous ventral projection, clava about 
twice as long as broad and without any conspicu- 
ous differentiated sensory areas or sensilla; fron- 
tovertex about one-third head width and with 
distinct piliferous punctures which descend part 
way between eyes and facial impression, facial 
impression remote from eyes; distance from 
anterior ocellus to facial impression about equal 
to width of frontovertex; forewing with lower 
part of apical margin straight. 

Material examined. 

COSTA RICA, lcf , San Jose, Braullio Carrillo 
NP, Est. Carrillo, 600m, 13. ii.1991 (Hym. 
Parataxonomists); 4cf, Limon, RB Hitoy- 
Cerere, 100m, 14-19.U991 (J.S. Noyes); lcf, 
PN Cahuita, 5m, l.iii.1991 (J.S. Noyes). Mate- 
rial in BMNH, INBio. 

Aenasius sp. B 

Diagnosis. Length about 1.33 mm. Forewings 
infuscate with a naked, hyaline streak at apex of 
venation; tarsi yellow, mid tibial spur dark 
brown; funicle 6-segmented, pedicel longer and 
wider than Fl, funicle segments subquadrate and 
without a conspicuous ventral projection, F6 and 
clava and without any conspicuous differentiated 
sensory areas or sensilla and clava about 2.5 x as 
long as broad; frontovertex about one-third head 
width and with distinct piliferous punctures 
which descend part way between the facial 
impression and eyes, facial impression remote 
from eyes; distance from anterior ocellus to facial 
impression about equal to minimum width of 
frontovertex. 

Material examined. 

COSTA RICA, lcf, Limon, RB Hitoy-Cerere, 

14-19.L1991 (J.S. Noyes). Material in BMNH. 



Notes on males 

In addition to the species treated above, we have 
recognised a further 20 species from males only. 
In order to facilitate their future recognition we 
provide, below, short diagnoses of the males of 
these species. 



Aenasius sp. C 

(Figs 68, 73) 

Diagnosis. Length 0.83-1.40 mm. Forewings 
hyaline (Fig. 68), sometimes evenly slightly 
infuscate; tarsi dark brown, mid tibial spur dark 
brown; scape widest below middle, funicle (Fig. 



ENCYRTIDAE OF COSTA RICA 



141 



73) 6-segmented, pedicel shorter and narrower 
than Fl which is trapezoidal in profile and up to 2 
x as long as F2, F2-F6 subquadrate but appear- 
ing slightly oblique in profile and with a con- 
spicuous ventral projection, F6 and clava without 
any conspicuous differentiated sensory areas or 
sensilla and clava nearly 2.5 x as long as broad; 
frontovertex about 0.4 x head width or less and 
with distinct piliferous punctures which do not 
extend between facial impression and eyes, facial 
impression touching eyes; distance from anterior 
ocellus to facial impression about 0.5-0.6 x 
minimum width of frontovertex. 

Material examined. 

COSTA RICA, 52d\ Guanacaste Prov., Santa 
Rosa NP, various localities, 1985-1989 (D. Jan- 
zen, I.D. Gauld); 20", Guanacaste Prov., Gua- 
nacaste NP, iii,1990 (J.S. Noyes); Id", San Jose, 
Ciudad Colon, 800m, iii-iv. 1990 (L. Fournier); 
lcf, San Jose, Ciudad Colon, Hda El Rodeo, 
800m, 16. ii. 1991 (J.S. Noyes). Material in 
BMNH, INBio. 

Aenasius sp. D 

(Figs 66, 74) 

Diagnosis. Length 0.73-0.88 mm. Forewings 
hyaline (Fig. 66); fore and mid tarsi yellow, mid 
tibial spur white, hind tarsi testaceous; funicle 
(Fig. 74) 6-segmented, pedicel shorter and nar- 
rower than Fl, funicle segments subquadrate and 
without a conspicuous ventral projection, F6 and 
clava with any conspicuous differentiated sensory 
areas or sensilla and about 2.5 x as long as 
broad; frontovertex about 0.4 x head width and 
with distinct piliferous punctures which descend 
part way between facial impression and eyes, 
facial impression remote from eyes; distance 
from anterior ocellus to facial impression about 
0.4 x minimum width of frontovertex. 

Material examined. 

COSTA RICA, 17cf, Guanacaste Prov., Santa 
Rosa NP, various dates 1986-1991 (D. Janzen, 
I.D. Gauld). Material in BMNH, INBio. 

Aenasius sp. E 

(Figs 67, 75, 104) 

Diagnosis. Length 0.76-1.02 mm. Forewings 
hyaline (Fig. 67); tarsi yellow, mid tibial spur 
yellow; funicle (Fig. 75) 6-segmented, pedicel 
shorter and narrower than Fl which is quadrate, 
funicle segments subquadrate and without a con- 
spicuous ventral projection, F6 and clava (Fig. 
104) with an oval sensory area and about 3 x as 



long as broad; frontovertex about 0.4 x head 
width and with distinct piliferous punctures, 
facial impression touching eyes; distance from 
anterior ocellus to facial impression about 0.5 x 
minimum width of frontovertex. 

Material examined. 

COSTA RICA: 12o\ Guanacaste Prov., Santa 

Rosa NP, various dates 1985-1987 (D. Janzen, 

I.D. Gauld); lcf, Guanacaste Pv, Guanacaste 

NP, ll.iii. 1990 (J.S. Noyes). Material in BMNH, 

INBio. 

Aenasius sp. F 

Diagnosis. Length 0.63-0.92 mm. Forewings 
hyaline; tarsi yellow, mid tibial spur white; 
funicle 6-segmented, pedicel about as long as Fl 
but slightly narrower, Fl quadrate; funicle seg- 
ments disc-like and without a conspicuous ven- 
tral projection, clava without any conspicuously 
differentiated sensory areas or sensilla and about 
2.5 x as long as broad; frontovertex about one- 
half head width and with distinct piliferous punc- 
tures which descend part way between eyes and 
facial impression, facial impression remote from 
eyes; distance from anterior ocellus to facial 
impression about 0.5 x minimum width of fron- 
tovertex. 

Material examined. 

COSTA RICA: 5o\ Guanacaste Prov., Santa 
Rosa NP, various dates 1985-1986 (D. Janzen, 
I.D. Gauld); lo", Puntarenas, Manuel Antonio, 
23-28.viii.1986 (L. Masner). Material in BMNH. 

Aenasius sp. G 

(Fig. 103) 

Diagnosis. Length about 0.83 mm. Forewings 
hyaline; tarsi yellowish brown, mid tibial spur 
white; funicle 6-segmented, pedicel longer but 
narrower than Fl, funicle segments disc-like and 
without a conspicuous ventral projection, F6 and 
clava with conspicuous ventral sensory areas 
(Fig. 103), clava about twice as long as broad; 
frontovertex a little more than one-third head 
width with distinct piliferous punctures which do 
not descend between facial impression and eyes, 
facial impression nearly touching eyes; distance 
from anterior ocellus to facial impression about 
0.5 x minimum width of frontovertex. 

Material examined. 

COSTA RICA: 2d", Guanacaste Prov., Santa 
Rosa NP, Hacienda 1-0, 6-27. ix. 1986 (D. Jan- 
zen, I.D. Gauld). Material in BMNH. 



142 



J.S. NOYES ANDH. REN 



Aenasius sp. H 

(Fig. 105) 

Diagnosis. Length 0.76-0.89 mm. Forewings 
hyaline; tarsi yellow, mid tibial spur yellow or 
brown; funicle 6-segmented, pedicel longer than 
Fl which is quadrate and about as wide, funicle 
segments disc-like and without a conspicuous 
ventral projection, clava with one or two peg-like 
sensilla (Fig. 105) and about twice as long as 
broad; frontovertex a little more than one-third 
head width and with distinct piliferous punctures 
which descend part way between eyes and facial 
impression, but facial impression remote from 
eyes; distance from anterior ocellus to facial 
impression about 0.7 x minimum width of fron- 
tovertex. 

Material examined. 

COSTA RICA, 2cf, Guanacaste Prov., Santa 
Rosa NP, Bosq. Hum. 11-0, ll.v-l.vi.1985 (D. 
Janzen, I.D. Gauld); 2d", Guanacaste Prov., 
Guanacaste NP, 500m, below Pitilla, 6. hi. 1990 
(J.S. Noyes). Material in BMNH. 

Comments. This material may represent more 
than one species. 

Aenasius sp. I 

Diagnosis. Length about 0.7 mm. Forewings 
hyaline; tarsi yellow, mid tibial spur yellow; 
funicle 6-segmented, pedicel longer than Fl and 
about as wide, funicle segments disc-like and 
without a conspicuous ventral projection, clava 
with one oval sensory area and about twice as 
long as broad; frontovertex about one-half head 
width and without distinct piliferous punctures, 
facial impression remote from eyes; distance 
from anterior ocellus to facial impression about 
0.5 x minimum width of frontovertex. 

Material examined. 

COSTA RICA, 2d\ Guanacaste Prov., Guana- 
caste NP, near Playa Naranja, 11. hi. 1990 (J.S. 
Noyes). Material in BMNH. 

Aenasius sp. J 

Diagnosis. Length 0.72-0.79 mm. Forewings 
hyaline; tarsi yellow to yellowish brown, mid 
tibial spur yellow; funicle 6-segmented, pedicel 
longer than Fl and about as wide, funicle seg- 
ments disc-like and without a conspicuous ven- 
tral projection, F6 and clava without conspicuous 
sensory areas, clava about twice as long as broad; 
frontovertex a little more than one-third head 
width, with relatively deep, reticulate sculpture 



and without distinct piliferous punctures, facial 
impression separated from eyes; distance from 
anterior ocellus to facial impression about 0.65 x 
minimum width of frontovertex. 

Material examined. 

COSTA RICA, 2cf, Guanacaste Prov., Lomas 
Barbudal Biol. Res. 15 Km SW Bagaces, 
28. ii. 1990 (J.S. Noyes); 2d", Guanacaste Prov., 
Guanacaste NP, near HQ, 2-20.iii.1990 (J.S. 
Noyes). Material in BMNH. 

Comments. This material possibly represents 
two separate species since there are slight differ- 
ences in the structure and arrangement of the 
sensilla on F6 and clava in the specimens from 
either locality. 

Aenasius sp. K 

Diagnosis. Length 1.30 mm. Forewings hya- 
line; tarsi yellow, mid tibial spur brown; funicle 
3-segmented, Fl transverse, clava without a 
naked ventral ridge, about 7 x as long as broad 
and 2.5 x as long as scape; frontovertex nearly 
half head width and with distinct piliferous punc- 
tures, two or three rows descending some of the 
way between eyes and facial impression; distance 
from anterior ocellus to facial impression about 
0.4 x minimum width of frontovertex. 

Material examined. 

COSTA RICA, lcf, Manuel Antonio NP, 

26.viii.1986 (L. Masner). Material in BMNH. 

Comments. Possibly a male of Aenasius phen- 
acocci or related species. 

Aenasius sp. L 

Diagnosis. Length 1.06 mm. Forewings hya- 
line; tarsi yellow, mid tibial spur dark white; 
funicle 3-segmented, Fl transverse, clava without 
a naked ventral ridge, about 3 x as long as broad 
and about 1.5 x as long as scape; frontovertex 
about one-third head width, with piliferous punc- 
tures obscured by relatively deep reticulate 
sculpture; distance from anterior ocellus to facial 
impression about equal to minimum width of 
frontovertex. 

Material examined. 

COSTA RICA, lcf, Puntarenas Prov., Mon- 

teverde, 15-16. vii. 1986 (L. Masner). Material in 

BMNH. 

Comments. Possibly a male Blepyrus sp. 



ENCYRTIDAE OF COSTA RICA 



143 



Aenasius sp. M 

(Figs 80, 94) 

Diagnosis. Length 0.94-1.44 mm. Forewings 
hyaline; tarsi dark brown, mid tibial spur dark 
brown; scape widest beyond middle, funicle (Fig. 
80) 5-segmented, Fl transverse, clava without a 
naked ventral ridge, about 4.5 x as long as broad 
and about 2.5 x as long as scape; frontovertex 
(Fig. 94) a little more than one-third head width, 
piliferous punctures indistinct and obscured by 
relatively deep irregular sculpture; facial impres- 
sion remote from eyes; distance from anterior 
ocellus to facial impression about 0.7 x mini- 
mum width of frontovertex. 

Material examined. 

COSTA RICA, 60", Guanacaste Prov., Santa 
Rosa NP, various dates 1985-1986 (D. Janzen; 
I.D. Gauld); Id". Limon, RB Hitoy-Cerere, 
100m, 14—19.i. 1991 (J.S. Noyes); lcf, San Jose, 
Ciudad Colon, Hda El Rodeo, 16.ii.1991 (Hym. 
Parataxonomists). Material in BMNH, INBio. 

Comments. This male is very similar to that 
described for Aenasius rnaplei Compere and is 
possibly therefore the male of Aenasius caeruleus 
Brues. 

Aenasius sp. N 

(Figs 81, 95) 

Diagnosis. Length 0.83-1.33 mm. Forewings 
hyaline; tarsi yellow-brown or testaceous-brown, 
mid tibial spur dark brown; funicle (Fig. 81) 
3-segmented, Fl transverse, clava without a 
naked ventral ridge, about 7-9 x as long as 
broad and about 3 x as long as scape; frontover- 
tex (Fig. 95) about one-third head width, pilifer- 
ous punctures indistinct and obscured by 
relatively deep reticulate sculpture, facial impres- 
sion remote from eyes; distance from anterior 
ocellus to facial impression about 0.7 x mini- 
mum width of frontovertex. 

Material examined. 

COSTA RICA, 13d", Guanacaste Prov., Santa 
Rosa NP, various localities, various dates 1986 
(D. Janzen; I.D. Gauld); 2o", Guanacaste Prov., 
Guanacaste NP, near HQ, 2-10. iii. 1990 (J.S. 
Noyes). Material in BMNH, INBio. 

Aenasius sp. O 

(Fig. 78) 

Diagnosis. Length 1.02-.27 mm. Forewings 
hyaline; tarsi yellow, mid tibial spur yellow; 



funicle (Fig. 78) 2-segmented, Fl transverse, 
clava with a naked ventral ridge, about 3 x as 
long as broad and about 2 x as long as scape; 
frontovertex a little less than one-third head 
width and with piliferous punctures obscured by 
shallow reticulate sculpture, facial impression 
remote from eyes; distance from anterior ocellus 
to facial impression about equal to minimum 
width of frontovertex or less. 

Material examined. 

COSTA RICA, 7d\ Guanacaste Prov., Santa 
Rosa NP, various localities, various dates, 
1985-1986 (D. Janzen, I.D. Gauld). Material in 
BMNH, INBio. 

Comments. This material probably represents 
males of A. insularis or a related species. 



Aenasius sp. P 

(Figs 79, 85) 

Diagnosis. Length 0.76-0.95 mm. Forewings 
hyaline; tarsi yellow, mid tibial spur white; 
funicle (Fig. 79) 2-segmented, Fl transverse, 
clava without a naked ventral ridge, about 6 x as 
long as broad and about 3 x as long as scape; 
frontovertex about one-third head width and 
with piliferous punctures obscured by shallow 
reticulate sculpture, facial impression remote 
from eyes; distance from anterior ocellus to facial 
impression about 0.75 x minimum width of 
frontovertex; genitalia as in Fig. 85. 

Material examined. 

COSTA RICA, 5a", Guanacaste Prov., Santa 

Rosa NP, various localities and dates 1985-1987 

(D. Janzen; I.D. Gauld). Material in BMNH, 

INBio. 



Aenasius sp. Q 

Diagnosis. Length 1.33 mm. Forewings infus- 
cate; tarsi yellow, mid tibial spur dark brown; 
scape widest in middle, funicle 2-segmented, Fl 
transverse, clava with a naked ventral ridge, 
about 4 x as long as broad and about 2.5 x as 
long as scape; frontovertex a little wider than 
one-third head width and with shallow piliferous 
punctures, one or two rows descending part way 
between facial impression and eyes; facial 
impression remote from eyes; distance from 
anterior ocellus to facial impression about 0.6 x 
minimum width of frontovertex. 



144 



J.S. NOYES ANDH. REN 



Material examined. 

COSTA RICA, ld\ Puntarenas Prov., Mon- 
teverde, St Luis Valley, 17.viii. 1986 (L. Masner). 
Material in BMNH. 

Aenasius sp. R 

(Fig. 82) 

Diagnosis. Length about 1.85 mm. Forewings 
with basal two-thirds infuscate; tarsi yellow, mid 
tibial spur dark brown; scape widest beyond 
middle and about 2 x as long as broad, funicle 
(Fig. 82) 2-segmented, Fl transverse and triangu- 
lar in profile, clava with a naked ventral ridge, 
about 4 x as long as broad and about 2 x as long 
as scape; frontovertex a little less than half head 
width and with distinct piliferous punctures 
descending between eyes and facial impression, 
thus clearly separating facial impression from 
eyes; distance from anterior ocellus to facial 
impression about 0.3 x minimum width of fron- 
tovertex. 

Material examined. 

COSTA RICA, 2cf , San Jose, Hwy 2, km 93, 

83°45'W 9°36'W (Cerro de la Muerte), 3200m, 

1-7. iv. 1985 (H. Goulet, L. Masner). Material in 

BMNH. 

Aenasius sp. S 

Diagnosis. Length about 1 mm. Forewings 
infuscate; tibiae with apices yellow, tarsi yellow, 
mid tibial spur yellow; scape widest below 
middle, funicle 2-segmented, Fl subquadrate, 
clava without a naked ventral ridge, about twice 
as long as broad and about as long as scape; 
frontovertex about one-third head width and 
with shallow piliferous punctures which extend 
between eyes and facial impression, facial 
impression remote from eyes; distance from 
anterior ocellus to facial impression about equal 
to minimum width of frontovertex. 

Material examined. 

COSTA RICA, 2cf, Puntarenas Prov., Mon- 

teverde, 15-16. vii. 1986 (L. Masner). Material in 

BMNH. 

Comments. This material may belong to the 
genus Blepyrus. 

Aenasius sp. T 

(Fig. 83) 

Diagnosis. Length about 0.9 mm. Forewings 
weakly infuscate; tarsi yellow, mid tibial spur 
brown; funicle (Fig. 83) 2-segmented, Fl trans- 



verse and triangular in profile, clava without a 
naked ventral ridge, about 5 x as long as broad 
and about 2 x as long as scape; frontovertex 
about 0.4 x head width and with inconspicuous 
piliferous punctures which extend between eyes 
and facial impression, facial impression remote 
from eyes; distance from anterior ocellus to facial 
impression about 0.7 x minimum width of fron- 
tovertex. 

Material examined. 

COSTA RICA, 2cf, Puntarenas Prov., Mon- 
teverde, St Luis Valley, 17.viii. 1986 (L. Masner). 
Material in BMNH. 



References 



Bennett, F.D. 1957. Trinidad Encyrtidae II. Some additional 
mealybug parasites. Canadian Entomologist 89: 569-572. 

Bruis. C.T. 1910. Some parasitic Hymenoptera of Vera Cruz, 
Mexico. Bulletin of the American Museum of Natural History 
28: 79-85. 

1921. A new Blepyrus. Zoologica, New York 3: 229-230. 

Clausen, C.P. (Editor) 1978. Introduced parasites and preda- 
tors of insect pests and weeds: A world review. United States 
Department of Agriculture, Agriculture Handbook 480: i-vi. 
1-545. 

Compere, H. 1931. New encyrtid (Hymenoptera) parasites of 
Pseudococcus species from Eritrea. University of California 
Publications in Entomology 5: 265-274. 

1937. The species of Aenasius. encyrtid parasites of 

mealybugs. Proceedings of the Hawaiian Entomological Soci- 
ety 9: 383-404. 

DeBach, P. & Warner, S.C. 1969. Importation and coloniza- 
tion of natural enemies of the striped mealybug, Ferrisia 
virgata (Hemiptera, Homoptera. Pseudococcidae) in Califor- 
nia. Annals of the Entomological Society of America 62: 
1117-1119. 

De Santis, L. 1980. Catalogo de los Himenopteros Brasilehos de 
laserie Parasitica incluyendo Bethyloidea. 395 pp. Editora da 
Universidade Federal do Parana. Curitiba. 

Flanders, S.E. 1940. A Brazilian parasite of Pseudococcus 
maritimus (Ehrh.). Journal of Economic Entomology 33: 
201. 

Funasaki, G.Y., Lai, P.Y., Nakahara, L.M., Beardsley, J.W. 
& Ota, A.K. 1988. A review of biological control introduc- 
tions in Hawaii: 1890-1985. Proceedings of the Hawaiian 
Entomological Society 28: 105-160. 

Gauld, I.D. 1986. Latitudinal species gradients in ichneumonid 
species-richness in Australia. Ecological Entomology 11: 
155-161. 

Gauld, I.D., Gaston, K.J. & Janzen, D.H. 1992. Plant alle- 
lochemicals. tritrophic interactions and the anomalous diver- 
sity of tropical parasitoids: the 'nasty' host hypothesis. Oikos 
65: 353-356. 

Gordh, G. 1979 Family Encyrtidae. In: Krombein, K.V., 
Hurd, P.D. jr. Smith, D.R. & Burks. B.D. (Editors) Catalog 
of Hymenoptera in America North of Mexico 1: 890-967. 
Washington, D.C. 

Gordh, G. & Hall, J.C. 1979. A critical point drier used as a 
method of mounting insects from alcohol. Entomological 
News 90: 57-59. 

Hayat, M. 1981. Taxonomic notes on Indian Encyrtidae 



ENCYRTIDAE OF COSTA RICA 



145 



(Hym.: Chalcidoidea). II. Journal of Natural History 15: 
17-29. 

Hayat, M., Alam, M. & Agarwal, M.M. 1975. Taxonomic 
survey of encyrtid parasites (Hymenoptera: Encyrtidae) in 
India. Aligarh Muslim University Publications (Zoological 
Series) on Indian Insect Types 9: i-iii, 1-112. 

Kaul, K.& Agarwal, M.M. 1985. Taxonomic studies on 
encyrtid parasitoids (Hymenoptera: Chalcidoidea) of India 
(with XVII plates). Aligarh Muslim University Publications 
(Zoological Series) on Indian Insect Types 13: i-v, 1-89. 

Kerrich, G.J. 1953. Report on Encyrtidae associated with 
mealybugs on cacao in Trinidad, and on some other species 
related thereto. Bulletin of Entomological Research 44: 
789-810. 

1967. On the classification of the anagyrine Encyrtidae. 

with a revision of some of the genera (Hymenoptera: Chalci- 
doidea). Bulletin of the British Museum (Natural History) 
(Entomology) 20: 143-250. 

Lbhr, B., Varela, A.M. & Santos, B. 1990. Exploration for 
natural enemies of the cassava mealybug, Phenococcus mani- 
hoti (Homoptera: Pseudococcidae), in South America for 
the biological control of this introduced pest in Africa. 
Bulletin of Entomological Research 80: 417-425. 

Maple, J.D. 1947. The eggs and first instar larvae of Encyrtidae 
and their morphological adaptations for respiration. Univer- 
sity of California Publications in Entomology 8: 25-122. 

Mercef, R.G. 1926. Los generos Chalcaspis How. y Eugahania 
nuevo (Hym. Chalc). Eos 2: 43-48. 

1947. Cuatro especics e un genero nuevo de Calcididos. 

Revista de la Real Academia de Ciencas, de Madrid 41: 
461-468. 

Noyes, J.S. 1980. A review of the genera of Neotropical 
Encyrtidae (Hymenoptera: Chalcidoidea). Bulletin of the 
British Museum (Natural History) (Entomology) 41: 
107-253. 



1989. The diversity of Hymenoptera in the tropics with 

special reference to Parasitica in Sulawesi. Ecological Ento- 
mology 14: 197-207. 

Noyes, J.S. & Hayat, M. 1994. Oriental mealybug parasitoids of 
the Anagyrini (Hymenoptera: Encyrtidae). viii + 554pp. 
CAB International. Wallingford, UK. 

Owen, D.F. & Owen, J. 1974. Species diversity in temperate 
and tropical Ichneumonidae. Nature 249: 583-584. 

Prinsloo, G.L. 1988. A review of the encyrtid wasp tribe 
Aenasiini, with descriptions of new Afrotropical taxa 
(Hymenoptera: Chalcidoidea). Journal of Natural History 
22: 1465-1482. 

Prinsloo, G.L. & Annecke, D.P. 1979. A key to the genera of 
Encyrtidae from the Ethiopian region, with descripitions of 
three new genera (Hymenoptera: Chalcidoidea). Journal of 
the Entomological Society of Southern Africa 42: 349-382. 

Shafee, S.A., Alam, M. & Agarwal, M.M. 1975. Taxonomic 
survey of encyrtid parasites (Hymenoptera: Encyrtidae) in 
India. Aligarh Muslim University Publications (Zoological 
Series) on Indian Insect Types 10: i-iii, 1-125. 

Swezey, O.H., Fullaway, D.T. & Sakimura, K. 1939. Recent 
records of the introduction of beneficial insects into the 
Hawaiian islands. Proceedings of the Hawaiian Entomologi- 
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Townes, H. 1972. A light-weight Malaise trap. Entomological 
News 83: 239-247. 

Walker, F. 1846. Characters of some undescribed species of 
Chalciditcs. (Continued) Annals and Magazine of Natural 
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Williams, D.J., Cox, J.M. & Yaseen, M. 1981. The cassava 
mealybug and its parasites in Paraguay and Bolivia. Biocon- 
trol News and Information 2: 88. 



146 



J.S. NOYES AND H. REN 









Figs 2-7 2, A. dives, antenna, $; 3, A caeruleus, antenna, J; 4, A. bolowi, antenna, $; 5, A regularis, 
antenna, $; 6, A. bolowi, scape, $; 7, A hyettus, scape, $. 



ENCYRTIDAE OF COSTA RICA 



147 









Figs 8-13 8, A. phenacocci, antenna, $?; 9, A. phenacocci, $, clava enlarged, showing longitudinal division of 
sensory area, $; 10, A. frontalis, antenna, $; 11, A frontalis, clava enlarged, $, showing lonitudinal division of 
sensory area; 12, A advena, antenna, <j>; 13, A insularis, antenna, $. 



148 



J.S. NOYES AND H. REN 



w 







14 








'f/'f; 




16 




17 








18 



19 



Figs 14-19 14, A. pelops, antenna, $; 15, A. paulistus, antenna, $>; 16, A. brasiliensis, antenna, 9; 17, A. 
mitchellae, antenna, $; 18, A. longiscapus, antenna, dark form, $; 19, A. longiscapus, antenna, pale form, $. 



ENCYRTIDAE OF COSTA RICA 



149 




21 










Figs 20-25 20, A. dives, forewing, ?; 21, A dives, venation enlarged, $; 22, A vexans, forewing, $; 23, /I. 
vejraw, venation enlarged, $; 24, A phenacocci, forewing, $; 25, /I. phenacocci, venation enlarged, $'. 



150 



J.S. NOYES ANDH. REN 



26 




27 









28 






31 








->yj/y j 



-v \\ Up hXxxn^ 



Figs 26-31 26, ,4. insularis, forewing, 9; 27, /I. insularis, venation enlarged, $; 28, /4. pelops, forewing, $: 29, 
A pelops, venation enlarged, 9; 30, A paulistus, forewing, $\3\,A. paulistus, venation enlarged, $. 



ENCYRTIDAE OF COSTA RICA 



151 




33 



■*** 




S# 












'fit, 












_j§ 



m 



L A 






Figs 32-37 32, A. frontalis, forewing, $ ; 33, A. frontalis, venation, enlarged, $ ; 34, A.bolowi, forewing, 9 ; 35, 
A. bolowi, venation, enlarged, 9; 36, A. hyettus, forewing, 9; 37, A. regularis, venation, enlarged, £. 



152 



J.S. NOYES AND H. REN 





40 














il%. 



Figs 38-43 38, A. caeruleus, forewing, ty\ 39, A. caeruleus, venation, enlarged, $; 40, A. brasiliensis , forewing, 
$; 41, A. brasiliensis, venation, enlarged, <J; 42, A mitchellae, forewing, 2; 43, /I. mitchellae , venation, 
enlarged, 2- 



ENCYRTIDAE OF COSTA RICA 



153 











44 


,,e': 




o". 1 ".'-'.'' ' 


#i 








^-^ 






, I 













■ ■■■':', 




-=£=>±\ \ 



46 






r 








Figs 44-49 44, A. longiscapus, forewing, £; 45, A. longiscapus, base of forewing, enlarged, $>; 46, /4. dives, 
apical gastral tergites, $; 47, /I. wexaws, apical gastral tergites, 9; 48, A. phenacocci, apical gastral tergites, $; 
49, /4. insularis, apical gastrs>l tergites, $. 



154 



J.S. NOYES AND H. REN 



I 

WA , 



50 



w^"^ 


: 


^0 


/ 




52 







Figs 50-55 50, Aenasius vexans, ovipositor; 51, A. vexans, proximal part of ovipositor, enlarged; 52, A. dives, 
ovipositor; 53, A. dives, proximal part of ovipositor, enlarged; 54, A. insularis, proximal part of ovipositor, 
enlarged; 55, A. phenacocci, ovipositor. 



ENCYRTIDAE OF COSTA RICA 



155 









Figs 56-61 Proximal part of ovipositor, photographed using phase contrast 
insularis; 59, A. mitchellae- 60, A. paulistus; 61, A. frontalis. 



56, A. dives; 57, A. pelops\ 58, A. 



156 



J.S. NOYES ANDH. REN 











1 Wffia 



66 




Figs 62-67 Forewings, cf - 62, /I. vejca/w; 63, /I. phenacocci; 64, A bolowi; 65, A brasiliensis; 66, sp. D; 67, sp. 



ENCYRTIDAE OF COSTA RICA 



157 









Figs 68-73 68, sp. C, forewing, O"; antennae, cf - 69, A. brasiliensis; 70, A. vexans; 71, A. bolowi; 72, sp. A 
(clava collapsed); 73, sp. C. 



158 



J.S. NOYES ANDH. REN 






^K 




\ 



78 




Figs 74-79 Antennae, d" - 74, sp. D; 75, sp. E; 76, A. paulistus, 11, A. advena; 78, sp. O; 79, sp. P. 



ENCYRTIDAE OF COSTA RICA 



159 







■t 



84 




Figs 80-85 80-83, antennae, cf - 80, sp. M; 81, sp. N; 82, sp. R; 83, sp. T; 84, A. advena, genitalia, cf ; 85, sp. 
P, genitalia, cf (taken using phase contrast). 



160 



J.S. NOYES ANDH. REN 









Figs 86-91 Head, facial view, $ - 86, A. vexans; 87, A. frontalis; 88, A. bolowi\ 89, A. caeruleus, 90, A. 
brasiliensis; 91, A. longiscapus . 



ENCYRTIDAE OF COSTA RICA 



161 








Figs 92-97 92, A. bolowi, clava, enlarged showing longitudinal division of sensory area, $; 93, A. caeruleus, 
clava, enlarged showing longitudinal division of sensory area, $; head, facial view, O" - 94. sp. M; 95, sp. N; 
A. bolowi, 97, A. advena. 



96, 



162 



J.S. NOYES ANDH. REN 




99 






101 







Figs 98-105 98, A. vexans, antenna (including reverse of clava), $; 99, A. vexans, genitalia, cf ; 100, A. pelops, 
right mandible, $; 101, A. vexans, right mandible, $; 102, A. bolowi, antenna, showing sensory patches on 
clava enlarged, cf ; 103, sp. G, F5, F6 and clava showing modified areas, cf (drawing of badly collapsed antenna 
and slightly modified); 104, sp. E, antenna, showing modified areas of F6 and clava enlarged, cf; 105, sp. H, 
antenna, showing modified area on clava enlarged, cf • 



ENCYRTIDAE OF COSTA RICA 



163 



INDEX 



Synonyms and misidentifications arc in italics; main citations in bold 



Acalypha 126 

acuminatus (Aenasius) 139. 140 

advena (Aenasius) 120, 121, 123, 126, 

127-128, 129-130, 139, 140, 147, 158, 

159, 161 
aesculifolia (Manihot) 135 
Albizzia 128 
Alchornca 134 
aliodera (Cordia) 137 
Aztcca 137 

Blcpyrus 136, 142, 144 

bolowi (Aenasius) 122, 133-135. 136. 

146. 151, 156. 157. 160. 161, 162 
Bougainvillea 132, 137 
brasilicnsis (Aenasius, Chalcaspis) 122, 

136-138, 148, 152, 156, 157, 160 
brevipcs (Dysmicoccus, Pseudococcus) 

120. 137, 139 

caeruleus (Aenasius) 122, 134-135, 
135-136. 143, 146. 152, 160, 161 

calceolariae (Pseudococcus) 132 

Carapa 140 

cariocus (Aenasius) 120, 122, 136, 137, 
138 

Catacnococcus 137 

Cccropia 137 

Chalcaspis 136 

cirrha (Aenasius) 121, 129. 130 

citri (Planococcus) 127 

ctavus (Pseudanasius) 119. 127 

colombiensis (Aenasius) 136, 137 

Conostegia 134 

Cordia 126. 137 

costaricensis (Alchornca) 134 

Crcmatogaster 137 

curassavica 126 

curvispinosa (Crcmatogaster) 137 

dives (Aenasius) 121. 123-124. 146. 149. 

153, 154. 155 
Dysmicoccus 120. 137. 139 

Ferrisia 120, 126, 127, 128, 133, 135. 
136, 139, 140 



flaiulersi (Aenasius) 125, 126 
frugilis (Pseudococcus) 132 
frontalis (Aenasius) 121, 133, 147, 151. 
155. 160 

gahani (Pseudococcus) 132 
Gliricidia 127. 135, 136, 139. 140 
gossypii (Phenacoccus) 126 
grenudensis 126 

Hedcra 132 
helix (Hedcra) 132 
hcrrcni (Phenacoccus) 120, 125. 126 
hibisci (Phenacoccus) 125 
Hibiscus 125. 126. 135 
byettus (Aenasius) 119. 126. 134. 136, 
146, 151 

ianihinus (Aenasius) 127. 128 
insignis (Cccropia) 137 
insularis (Aenasius) 121, 129. 130, 
132-133, 143. 147, 150, 153. 154, 155 

kerrichi (Aenasius) 121. 130-131. 132 

Lantana 125, 126 
lebbeck (Albizzia) 128 

longiscapus (Aenasius) 122, 123, 138. 

139-140, 148, 153. 160 
longispinus (Pseudococcus) 127, 132 
lua (Aenasius) 121, 128-129 

Macquilla 127 

madeirensis (Phenacoccus) 126 

Manihot 135 

manihoti (Phenacoccus) 120. 125 

maplei (Aenasius) 119. 135. 136. 143 

maritimus (Pseudococcus) 120. 132 

martinii (Neodiscodes) I 19 

mitchcllac (Aenasius) 122. 138-139. 148. 

152, 155 
montividensis 125, 126 

Neodiscodes 119. 120 
neobrevipes (Pseudococcus) 137 
nervilolia (Thcrvctia) 127 



oleae (Saissetia) 127 

pacificus (Aenasius) 139. 140 
paulistus (Aenasius) 120, 121, 123. 129. 

130. 132. 132-133. 148. 150. 155, 158 
pelops (Aenasius) 121, 129, 131-132, 

148. 150, 155. 162 
personatus (Aenasius) 135. 136 
phenacocci (Aenasius) 120. 121, 123, 

125-127. 128. 139. 142. 147. 149. 153, 

154, 156 
Phenacoccus 120, 125. 126. 132. 137 
Piper 140 
Pittosporum 126 
Planococcus 127 
Pseudanasius 119. 127 
Pseudococcus 120. 127. 128. 132. 137 

regularis (Aenasius) 136. 146. 151 

Saissetia 127 

saman (Samanea) 128 

Samanea 128 

Sida 126 

similis (Aenasius) 133. 134 

sociabilis (Pseudococcus) 132 

tachigaliae (Blcpyrus) 135 
theobromue (Aenasius) 136. 137 
Thcrvctia 127 
tUiaceus (Hibiscus) 135 
tomentosa 127 
trifolia (Virex) 128 
Tussacia 126 

vadosus (Aenasius) 139. 140 
vexans (Aenasius) 120. 121. 122. 

124-125. 126-127. 149. 153. 154, 156. 

157. 160. 162 
Virex 128 
virgata (Ferrisia. Pseudococcus) 120. 

126. 127. 128. 133. 135, 136, 139. 140 

xalepeh (Conostegia) 134 
xanthochora (Aztcca) 137 



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P. C. "Barnard. 1990. Pp. 117-286. 

Volume 60 

No. 1 The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae. Bombini). P.H. Williams. 

1991. Pp. 1-204. 

No. 2 Sattleria: a European genus of brachvpterous alpine moths (Lepidoptera: Gelechiidae). L.M. 
Pitkin & K. Sattler. 1991. Pp. 205-241. 
A review of wing reduction in Lepidoptera. K. Sattler. 1991. Pp. 243-2SS. 

Volume 61 

No. 1 Thrips (Thysanoptera) from Pakistan to the Pacific: a review. J.M. Palmer. 1992. Pp. 1-76. 

No. 2 Neotropical red-brown Ennominae in the genera Tkysanopyga Hcrrich-Scaffcr and Perissopteryx 
Warren (Lepidoptera: Cieometridae). M. Kruger&M.J. Scoble. 1992. Pp. 77-148. 

Volume 62 

No. 1 Caloptilia leal-miner moths (Gracillanidae) of South-East Asia. Decheng Yuan and Gaden S. 

Robinson. 1993. Pp. 1-37. 
No. 2 Neotropical Emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular 

reference to the species of Costa Rica (Lepidoptera: Geometridae. Geometrinae). Linda M. 

Pitkin. 1993. Pp. 39-159. 

Volume 63 

No. 1 A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: Encyrtidae). parasitoids of 
the immature stages of economically important insect species (mainly Hemiptera and 
Lepidoptera). D.W. Huang and J. S. Noyes. Pp. 1-135. 

No. 2 A taxonomic review of the common green lacewing genus Chrysoperla (Neuroptera: 
Chrysopidae). S.J. Brooks. Pp. 137-210. 

Volume 64 

No. 1 Revision of the neotropical genus Oospila Warren (Lepidoptera: Geometridae) M. A. Cook and 
M.J. Scoble 







117 Encyrtidae of Costa Rica (Hymenoptera: Chalcidoidea): the genus 
Aenasius Walker, parasitoids of mealybugs (Homoptera: 
Pseudococcidae) 

J.S. Noyes and H. Ren 



ENTOMOLOGY SERIES 

Vol. 64, No. 2, November 1995